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DEPARTMENT OF COMMERCE 

BUREAU OF FISHERIES 

HUGH M. SMITH, Commissioner 



THE MUSSELS OF THE CUMBERLAND RIVER 
AND ITS TRIBUTARIES 

By Charles B* Wilson and H* Walton Clark 



Bureau of Fisheries Document No, 781 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1914 



-si-un 



D. OF D. 
FEB 



« 



THE MUSSELS OF THE CUMBERLAND RIVER 
AND ITS TRU3UTARIES 

By Charles B. Wilson and H. Walton Clark 



Bureau of Fisheries Doccment No. 781 



CONTENTS. 



Page. 

Introductory 3 

The Cumberland River 4 

General description 4 

Physiography 5 

Comparison with the Maumee and Kankakee Rivers 6 

Characteristics of the mussel fauna 6 

Geographical distribution 7 

Contrast between the river above and below the falls 7 

Faunistic divisions below the falls 8 

First section — Cumberland Falls to Celina, Tenn 8 

Second section — Celina to Nashville, Tenn 9 

Third section — Nashville to Dover, Tenn 11 

Fourth section — Dover, Tenn. , to Smithland, Ky 12 

Tabular statement of distribution of species 13 

Relative abundance of different species 20 

Summary of distribution 21 

Notes on various stations 23 

Upper river and its tributaries 23 

River below the falls and its tributaries 24 

Character of water of the Cumberland River 38 

Commercial value of Cumberland shells 39 

Breeding season of Cumberland mussels 41 

Pearls and pearling 42 

Discussion of mussel species 45 

2 



Digitized by the Internet Archive 
in 2011 with funding from 
The Library of Congress 



http://www.archive.org/details/musselsofcumberlOOwils 



U. S. B. F— Doc. 781. 



Plate I. 






FIG. 1.— TRUNCILLA WALKERI, NEW SPECIES. 
Upper figures, females; lower figures, males. 




FIG. 2.— THE GREAT FALLS OF THE CUMBERLAND, 85 FEET HIGH, A BARRIER TO THE ASCENT OF 

FISH AND MUSSELS. 



THE MUSSELS OF THE CUMBERLAND RTVER AND ITS 

TRIBUTARIES. 



By Charles B. Wilson and H. Walton Clark. 



INTRODUCTORY. 



The purpose of this investigation was to ascertain the distribution, 
relative abundance, and habits of the various mussel species living 
in the river and its tributaries, and to make an intelligent appraisal of 
the mussel resources of the river from a commercial standpoint. 

The party was under the supervision of Dr. Robert E. Coker, 
director of the United States Biological Station at Fairport, Iowa, who 
furnished general instructions to be used by all field parties engaged 
in mussel investigations. In addition to the authors, the party 
included the late Mr. J. F. Boepple, the shell expert of the Fairport 
station, and Mr. Ernest Danglade, now scientific assistant in the 
Bureau of Fisheries, each of whom contributed fully as much as either 
of the authors to the success of the investigations. 

The work was begun about May 10 near the mouth of the Cumber- 
land River, and conducted thence upstream through the State of 
Kentucky and into Tennessee as far as Clarksville. During the pre- 
vious year it had been carried from Pineville, Ky., to Celina, Tenn. 
Accordingly, it was now resumed at Celina, where the Obey River, 
a tributary of the Cumberland from the south, was investigated. 
Thence the work continued slowly down the Cumberland itself. 

From Jellico, Tenn., and Williamsburg, Savoy, Corbin, Livingston, 
and Barbourville, Ky., as centers, the upper portions of the Cumber- 
land River, the Clear Fork, Big South Fork, Laurel and Rock Castle 
Rivers were examined. Neither the main river nor any of these 
tributaries is navigable for a boat, so that the investigations had to 
be conducted by team, driving along the banks or visiting convenient 
fords and shallows. 

The party then drove by team from Williamsburg to the Cumber- 
land Falls, proceeded again by team from the falls to Parkers Lake 
station, and thence by rail to Burnside, Ky. This is the head of 
steamboat navigation on the river, and here a small boat was con- 
structed in which to proceed down the main river, thus completing 
the survey of the entire river. 

3 



4 MUSSELS OF CUMBERLAND EIVER AND TRIBUTARIES. 

During ail these investigations the methods followed by the two 
divisions of the party were made as different as possible in order to 
cover the field more thoroughly. Mr. Boepple used the crowfoot 
dredge, tongs, and mussel rake, and worked the deeper portions of the 
river. The rest of the party covered the shallower water, riffles, sand 
bars, and smaller tributaries, and, of course, obtained the mussels by 
wading. 

A careful record was kept of the temperature of the water at the 
various stations, and as often as seemed advisable samples were 
taken for subsequent analysis. 

In addition to making original observations the party secured as 
much information as could be obtained from local fishermen and 
clammers with reference to the location of the mussel beds, past and 
present operations upon them, and the finding of pearls and baroques. 

For such information we are particularly indebted to the following 
persons: Mr. Walter, of Dover, Tenn., an extensive dealer in shells; 
Mr. Samuel Dabbs, a clammer of Dover; Mr. M. K. Clark, proprietor 
of the blank factory at Clarksville, Tenn.; and Mr. Cicero Harris, a 
boatman who had floated down from the upper part of the river fishing 
and clamming, and who knew the river more intimately than anyone 
else it was our fortune to meet. To these gentlemen as well as to 
many others who extended favors and assistance whenever oppor- 
tunity offered, our sincere thanks are tendered. 

As fast as they were obtained, the samples of water and specimens 
were shipped to the biological station at Fairport. The shells were 
subsequently identified and studied by the principal author with the 
results herein set forth. 

THE CUMBERLAND RIVER. 
GENERAL DESCRIPTION. 

The main branch of the Cumberland River rises among the foot- 
hills of the Pine Mountains, in the southeastern corner of Kentucky. 
It flows southwest along the eastern side of the mountains, receiving 
many tributaries. Near Pineville it turns at a right angle and flows 
northwest through a wide gap in the mountains, and then swings to 
the south, its general course being that of a half circle, convex toward 
the north. At State Line, in Monroe County, it crosses into Tennes- 
see, its general course in the latter State being also that of a half 
circle but convex toward the south. 

At Tobaccoport, in Stewart County, it crosses the State line back 
into Kentucky, flows northwest and enters the Ohio at Smithland, 
only 12 miles above the mouth of the Tennessee River at Paducah. 
The distance from the source to the mouth in a straight line is about 
325 miles, but the river is so extremely crooked that its total length 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 5 

is nearly 750 miles. Its principal tributaries are the Laurel and Rock- 
castle Rivers from the north, which join it within a few miles of each 
other at the southwestern corner of Laurel County, Ky.; the Big 
South Fork, whose mouth is at Burnside, Ky. ; the Obey River from 
the south at Celina, Tenn.; Roaring River, from the south, at Gaines- 
boro Landing, Tenn.; Caney Fork, from the south, at Carthage, 
Tenn.; Stones River, from the south, 15 miles above Nashville 
Tenn.; Harpeth River, from the south, at Pardue, Tenn.; and the 
Red River, from the north, at Clarksville, Tenn. 

The Cumberland is navigable during high water from its mouth to 
Burnside, Ky., a distance of 525 miles, and a system of locks is in 
process of construction which will make navigation possible during 
the entire year. 

PHYSIOGRAPHY. 

The area drained by the river and its tributaries is about 25,000 
square miles, and embraces mountain ranges, a continental plateau 
(the Cumberland Plateau), and lowlands. Alorg the upper reaches 
of the river amorg the Cumberland and Pine Mountains in the eastern 
portion of the plateau the rocks are largely Cambrian sandstone; 
through the remainder of the plateau and the long stretch of lowlands 
they are almost universally limestone. The dividirg line is at 
Cumberland Falls in the western part of Whitley County, Ky., where 
the river plunges over a wall 85 feet in height. From the source to 
the falls the river has nowhere cut its channel very deep; below the 
falls, and especially through the plateau, the banks are lined almost 
continuously with high limestone cliffs, filled with caves and roughly 
weathered. The faces of these cliffs furnish abundant evidence of past 
upheavals in numerous faults and contortions of the strata, as well 
as in repeated anticlinal and synclinal folds, differing considerably 
in intensity at different localities. 

Above the falls the river valley is comparatively narrow, but below 
the falls it widens somewhat, and the river winds back and forth in 
broad and then in shorter curves, with cliffs now on one side and now 
on the other. 

So evenly has the channel been worn down through the soft lime- 
stone that there are no rapids of any importance below the falls, and 
steamboats can run from the mouth up to Burnside in Pulaski 
County, Ky., within comparatively few miles of the falls, as already 
stated. This makes the river easy to navigate for two-thirds of its 
entire length, and since it runs through a great region remarkable for 
its mineral and agricultural resources and its large forests, but with a 
physical contour which makes the building of railroads exceedingly 
expensive, the Cumberland is destined to be one of the most important 
commercial highways of the United States. 



6 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

COMPARISON WITH MAUMEE AND KANKAKEE RIVERS. 

Both the Maumee and Kankakee Rivers, which were examined by 
the present authors, are situated in regions profoundly modified by 
the great glacier. In their basins the ice mass first removed the 
entire fauna and flora, and when it melted established new channels 
by which the river was restocked. 

The Cumberland Valley presents an entirely different history. It 
is situated in a region which is geologically very old and which has 
not been much disturbed since its first upheaval, except by the ordi- 
nary forces of weathering and erosion and the subsequent formation 
of mountains. The Cumberland and Pine Mountains, as well as the 
great Cumberland Plateau, are portions of the Appalachian system, 
and the wrinklirg which formed them took place toward the close of 
the Upper Silurian period. Originally very much higher than at the 
present day, they have gradually yielded to weathering and erosion, 
but are otherwise unchanged. The great glacier reached only a little 
below the Ohio River, which is far to the north of the Cumberland 

Valley. 

CHARACTERISTICS OF THE MUSSEL FAUNA. 

Consequently a primitive fauna and flora are to be looked for in 
this valley, one that began with the very origin of the valley itself, 
and has been gradually developing ever since without any serious 
disturbance; and in fact the best American authorities regard the 
Mississippi Valley as the original home of fresh-water mussels upon 
this continent, the rest of the rivers, ponds, and streams having been 
populated from this source. Some authorities even say that there is 
evidence to show that this fauna developed first in the New World and 
then spread to the Old World. However that may be, it is certain 
that the Mississippi area has the greatest diversity of species and the 
most magnificent shells to be found anywhere in the world. 

The Cumberland and Tennessee Valleys are among the very oldest 
portions of the Mississippi region, and are commonly looked upon as 
the center of this wonderful mussel fauna. Accordingly we should 
expect to find in them a great diversity of species, some of which 
would be found nowhere else, and that such is the case has been well 
shown by many conchologists. Over 80 different forms of mussels 
have been reported from the Cumberland River, and the present 
examination has added 3 others. This is considerably more than 
twice the number found in the Maumee or the Kankakee River 
systems, and is a remarkably large representation compared with any 
river of equal size. A few of these species have never been reported 
from any other locality, but the great majority are common to the 
southern portion of the Mississippi system. Such of these as were 
found during the present examination are enumerated on pages 14 
to 19. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 7 

GEOGRAPHIC DISTRIBUTION OF THE MUSSELS. 
CONTRAST BETWEEN THE RIVER ABOVE AND BELOW THE FALLS. 

The Cumberland Falls establish a natural barrier, dividing the river 
into an upper one-third and a lower two-thirds, between which there 
can be practically no interchange of animal life, and very radical 
differences appear in the mussel fauna. Above the falls only a very 
few species of mussels are found, and these are considerably dwarfed. 
TJnio gibbosus is the only species in any abundance, and rarely one 
may find examples of Lampsilis ovata, Alasmidonta minor, and Ano- 
dontoides ferussaciana. This scarcity of species is as much due to the 
fact that all the conditions are unfavorable (see p. 23) as it is to the 
lack of intercourse past the falls, and in all probability there would 
be very little profit in stocking the river above the falls with mussels. 
Indeed we were told that some Lampsilis ovata were taken from below 
the falls and transplanted to the river above about seven years ago, 
with visible results, possibly, in the few dwarfed specimens of this 
mussel now present in the upper river. 

In the river below the falls conditions are totally different. In 
the very pool at the base of the falls were obtained 19 species of 
mussels, all of them of normal size and perfectly healthy. And from 
this point down to the Ohio every portion of the river bed that is at 
all suitable for mussels is fairly covered with them. 

Much of this part of the river has been thoroughly worked over 
by agents of the button factories, and the location, extent, and pos- 
sibilities of the various beds are well known. Some clammers even 
have a memorandum list of the beds, giving the percentages of usable 
and useless shells in each. Many of these beds have been worked 
for some time, a few of them as long as 10 years, and an immense 
number of shells have been taken, as many as 200 to 300 tons from 
some of them. But in spite of the great number of mussels taken 
out, the river as a whole, according to general accounts, does not 
show any marked depletion except in one or two restricted locali- 
ties. On the contrary, a comparison of many beds in the vicinity 
of Celina, Tenn., examined by Mr. Boepple in 1910 and again in 
1911, showed a considerable increase. This was especially true of 
beds situated above the silt in the back water from the various 
lock dams. Such places seem peculiarly suited to rapid mussel 
growth, and furnish thereby a valuable suggestion as to the best 
localities for artificial propagation. 

Of course the mussels that were too close to the dams, or that were 

in the mouth of tributaries filled with back water from the dams, 

would be killed by the increased deposit of silt, and the rise of 

water from behind the dams makes it harder to secure the mussels. 

4788°— 14 2 



8 MUSSELS OF CUMBERLAND EIVER AND TRIBUTARIES. 

On the whole, however, the benefits seem greater than the disad- 
vantages. 

Incidentally it is worthy of note that the water privileges at 
Cumberland Falls have been leased to a company which has already 
begun operations toward establishing a power plant for furnishing 
electricity to Louisville and other cities. 

FAUNISTIO DIVISIONS OF THE RIVER BELOW THE FALLS. 

For our present purpose we may divide the river below the falls 
into four sections, fairly well separated by natural conditions, and 
by differences in the relative numbers of the various mussels. These 
sections will be discussed in order, beginning at the falls and pro- 
ceeding toward the mouth of the river. 

First section, from Cumberland Falls to Celina, Tenn., 17 h miles. — ■ 
While there are numerous and rich mussel beds along this portion 
of the river, there is no commercial clamming. This is chiefly due 
to the high percentage of culls, small species, and pinks, the latter 
mostly elephant-ear (Unio crassidens). The most important com- 
mercial mussel is the southern mucket (Lampsilis ligamentina gibba). 

The elephant-ear is not killed in any great numbers by pearlers 
because it is not looked upon as a pearl-bearing species, while other 
mussels, supposed to contain pearls, are often nearly exterminated. 
Up to the present time, moreover, this mussel has been refused by 
the buyers for button factories. Consequently it has been neg- 
lected or culled out by the fishermen in the lower sections of the 
river and left comparatively free to breed, the glochidia to be picked 
up by fish and carried up toward the falls. Natural conditions have 
in some way also given the purple spike ( Unio gibbosus) an advantage 
over other species above the falls. Similar conditions may have 
been equally favorable to the closely related elephant-ear below the 
falls. Perhaps these considerations will help to explain their pre- 
ponderance in these two localities. 

There are 19 mussel beds in this section of the river and the pro- 
portion of commercial shells and culls, together with the size of 
the bed and the kind of bottom, are shown in the following table: 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 
Fikst Section, Cumberland Falls to Celina, Tenn. 





Percentage of commercial 
shells. 


Percentage 
of culls. 


Conditions. 


Mussel beds. 


0> 

M 
o 

3 


o5 
o 

s 


■S 

(3 
O 

<B 

be 
be 


"3 

■a 

a 

63 
CO 


a 

o 

1 
o 
o 


M 
o 

63 

u 

53 

is 


J* 
■ft 

CO 




.9 

°«5 
03 M 

a 

CO 


Size of bed. 


Kind of 
bottom. 


0) 

H 

I . 

& u 

a° 


9 
H 

®g 

ft? 

|o 

Eh 




24 
5 

7 

30 

35 






1 

4 
3 


6 
3 


10 
10 

10 

2 


15 
26 

29 

16 

16 

20 

1 

2 

20 

20 

6 

6 

1 


7 
15 

5 

28 
28 
60 
14 
20 
50 
45 

"40 
39 
50 
50 
40 
60 
18 
20 


25 

37 

40 

20 

10 

20 

40 

25 

10 

10 

70 

4 

20 

8 

6 

8 


Large 

...do 


Rocks 

...do 


e j? 
81 ' 

92 
86 


• F. 
84 


Big South Fork opposite Par- 






86 


kers Lake 

Big South Fork above Burn- 
side 

Railroad bridge, Burnside 






Medium... 
Small 


Gravel 

...do 


82 








Fishing Creek Bar 






5 




...do 


...do 


86 
86 
86 


85 


Fords Island 










Large 

Medium. . . 


Sand 

Gravel 

...do 


85 




5 
22 
10 
12 

4 
40 

5 
40 
40 
30 
35 
10 
10 


2 




1 


9 

9 


32 
15 


8* 






One mile below Lock 21 






2 
3 


...do 
...do 
..do 


Mud 

Sand 


86 


85 


Horseshoe Bottom 


















5 

5 

.... 

3 
6 


10 

2 
5 


...do 


80 
87 


81 




1 
9 
1 


'u 


2 
1 


...do 

Small . 


Gravel 

do... 


81 








Medium... 


...do 






Sells Shoals 










...do 








2 
1 

10 
10 


2 

1 


2 


8 


2 

1 
2 
4 


Medium... 
..do 








Champs Shoals 








Biggerstaff Bar 






33 
6 


15 
30 




...do 


80 


82 




2 




6 


do 


...do 















The table shows at a glance that the proportion of culls is so large 
in nearly every one of the beds that they yield but a poor profit to 
the clammer. 

The conditions, however, are everywhere favorable to mussel 
growth, as is evidenced by the number and variety of the shells. 
These mussel beds each contain a fair proportion of commercial 
shells, three of which, the southern mucket, the butterfly, and the 
Ohio River pigtoe, might well be propagated artificially. In this 
way the preponderance of culls could be greatly reduced in a few 
years, if not wholly overcome. 

Although there is no clamming, there is considerable pearling in 
this section of the river and large piles of shells were found in a number 
of places where the pearlers had left them. This was especially true 
at Fords Island, Mill Springs Bar, below Lock 21, Wells Island, 
Selfs Shoals, and Champs Shoals. It will be noticed that in coming 
down the river the first pigtoes were found at Mill Springs Bar and 
the second lot at Indian Creek Shoals. 

Second section, from Celina to Nashville, Tenn., 190 miles. — The 
mussel beds increase a little in number and considerably in size 
along this section of the river, and in consequence there is more 
commercial shelling. The percentage of pinks and spikes steadily 
decreases, especially that of the former, and there is a corresponding 
increase in the commercial species. The Ohio River pigtoe becomes 
the most common button shell, while the elephant-ear not ouly 
decreases in numbers, but partially changes its color, and with 



10 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



white nacre it answers fairly well for button making. The condi- 
tions are even better suited for mussel propagation than in the pre- 
ceding section. 

The following table gives the percentages of the various mussel 
species and other useful data: 

Second Section, Celina to Nashville, Tenn. 





Percentage of commercial 
shells. 


Percentage 
of culls. 


Conditions. 


Mussel beds. 


CD 

M 


o 

5 




a 

a 

03 
GO 


o 
o 

M 
o 
Ph 


% 


CJ 
>> 

53 

is 




ft 
02 


a 

s 


a 

■~ — 
03 m 

a 

02 


Size of bed. 


Kind of 
bottom. 


ID 

H 

a 

03 c 
fe'S 

ft-* 

1° 






25 

20 

5 

20 

6 

20 
20 
20 
10 
10 

3 

20 
30 
20 
20 


30 
25 


15 
10 


.... 


5 
10 


5 
10 


2 
5 


17 
20 





Small 
...do 


Gravel 


"F. 


°F. 




...do 








85 
5 

10 


...do 


...do 








30 

20 
15 
15 
25 
5 
40 
37 
35 
25 
15 
40 


10 

15 
15 
15 
10 
5 
10 
10 
10 
10 
10 
10 


1 
.... 

2 
1 


3 
5 
6 
5 


5 

20 
10 
15 
12 
5 
4 
18 
15 
10 
12 
10 


10 

"ft 7 

2 

"6" 
2 
2 
1 
5 
5 


20 

17 
20 
20 
25 
70 
20 

4 
15 

6 
12 

5 


Large 

Medium... 
...do 


Sand and 
mud. 

Gravel 

...do 








80 


79 








4 


Large 


...do 


80 


83 




SmaU 


...do 






Large 


...do 








2 

2 


10 
"2 


...do 


...do 








10 


...do 


...do 




82 




...do 


...do 


82 


85 




10 
10 


...do 


...do 






1 

2 


8 
5 


...do 


...do 


91 

84 


85 




...do 


...do 


85 













In addition to the beds above enumerated, small and not very 
profitable ones were reported by local clammers at Billiards Gap, 
8 miles below Simpsons Island; at Wartrace Creek Bar, 4 miles 
further down the river; at Pinks Bar, 2 miles below; at Lower 
Holliman Island, a mile below Phillips Branch; at the head of 
Sullivans Island, 5 miles lower; at the foot of the sand shoals near 
Haneys Landing; at Turkey Creek Shoals, just above Carthage; at 
Hunters Point, a mile below Lock No. 5; at the mouth of Spring 
Creek, 5 miles above Cairo; at the foot of Cunningham Island, 
2 miles nearer Cairo; at Mauskers Island, just above Edgefield Junc- 
tion; and at Priestly Shoals, 5 miles above Nashville. 

At Gainesboro Landing the mussels were all obtained from Roar- 
ing River, a tributary of the Cumberland from the south (see p. 29). 

At Cotton Bar 12 tons of shells were cribbed along the bank, of 
which 60 per cent were pigtoes; washboards, monkey-faces, and 
butterflies were also common. Simpsons Island was the highest 
point on the river where clammers were found actually at work. 

Muskrats were making heavy inroads into the mussel beds at several 
places, notably at Puryears Bar, at Mauskers Island, and Hills Island. 
All the piles of shells left by these animals showed that they have a 
decided preference for pigtoes. 



MUSSELS OF CUMBERLAND RIVER A1STD TRIBUTARIES. 



11 



Third section, from Nashville to Dover, Tenn., 105 miles. — This por- 
tion of the river has been more thoroughly worked by the clammers 
than has any other. It contains the largest and most valuable mussel 
beds of the entire river, and the location of all the beds, together with 
their size and relative value, are. well known. The proportion of 
merchantable shells, moreover, has increased until there is no longer 
any locality in this part of the river where the pinks and spikes pre- 
ponderate. The Ohio River pigtoe still continues to be the most 
common and valuable commercial shell, but the niggerhead becomes 
a close second and from Clarksville to Dover outranks the pigtoe. 

So much does the commercial clamming increase and so great is the 
influence of the ready local market for shells that pearling as a dis- 
tinctive vocation practically disappears. Every clammer is on the 
watch for such pearls as may be found in the shells which he cleans 
for the market, but there is very little hunting for pearls with no other 
object in view. This increase in the commercial clamming is due 
almost entirely to the activity of the button-blank factory at Clarks- 
ville, near the center of this third portion of the river, which furnishes 
a convenient market for all the shells taken in the vicinity. 

The proprietor of this factory, Mr. M. K. Clark, is much interested 
in everything that pertains to clamming, and with his assistance sev- 
eral thousand glochidia' of the yellow sand-shell were taken from ripe 
female mussels and placed in tubs of water with small fish caught in 
adjacent ponds. After the young mussels had fastened themselves 
to the fish the latter were turned loose in the river. This was the 
first time that mussels had ever been artificially planted in the 
Cumberland. Mr. Clark also gave us most of the data for the follow- 
ing: table of mussels beds: 



Third 


Section, Nashville 


TO 


Dover, Tenn. 










Percentage of commercial 
shells. 


Percentage 
of culls. 


Conditions. 


Mussel beds. 


o 


CD 
O 

£ 


(3. 

at 
43 

© 
to 
to 


"CD 

43 

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40 
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55 
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53 
74 
38 
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7 
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90 


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1 I 



12 MUSSELS OF CUMBERLAND RIVEE AND TRIBUTARIES. 

There are also small beds containing a limited number of market- 
able species at the following localities: Just below Lock No. 1, along 
the north bank of the river, badly depopulated by sand dredges ; near 
the Tennessee Central Railroad bridge, also along the north bank; at 
Whites Creek Bar, considerably dug up by sand dredging; along the 
mouth of Indian Creek, 20 miles below Nashville; below Lock A on 
the south bank of the river ; at Betsytown on a very rough and rocky 
bottom; at Davis Riffle extending diagonally across the river; oppo- 
site the pumping station of the Clarksville waterworks; at Kentucky 
Landing and Red Rock Landing, the latter bed nearly worked out; 
at Palmyra Island along the west bank of the river; at Cumberland 
City just below the steamboat landing; and at Wells Island, 2 miles 
farther down the river. 

Thus the third section of the river contains a larger number of 
mussel beds than any of the other sections, and the beds are richer 
both in numbers and species of mussels. It is the section of the pig- 
toe and niggerhead mussels, and those species are the most abundant 
button shells. There has also been a marked increase in the yellow 
sand-shell and the monkey-face. 

This portion of the river, however, is also the nearest to the center 
of demand, and consequently its beds have been worked longer and 
harder than any of the others. The most of them do not show any 
signs of depletion but remain as rich as when the work first began. 
The most important beds are, for the conchologist, the one at Half 
Pone Bar, where the smaller and rarer species are specially abundant, 
and for the button man the one at Guisers Bar, which has yielded 
rich returns through a long series of years; in fact, from the very 
beginning of work here on the river. 

Fourth section, Dover to Smiihland, Ky., 85 miles. — While this 
section is not as well known as the preceding, and has not been 
worked as much, it probably contains as many and as valuable 
mussels. 

The center of demand was still the blank factory at Clarksville, 
to which all the shells have to be transported up the river. But a 
sort of secondary center has been established at Dover, Tenn., where 
Mr. Walter, one of the leading merchants of the town, purchased 
most of the local shells and hired most of the clammers. Further- 
more, the business in this part of the river was conducted in the most 
approved and up-to-date manner. The boats were towed to and from 
the mussel beds by small launches, the mussels themselves were con 
veyed from the boats up the steep river bank by steam power, and 
were finally cleaned by steam conveyed to the pans in a pipe from the 
engine. 



MUSSELS OF CUMBERLAND EIVEE AND TRIBUTARIES. 
Fourth Section, Dover to the Ohio River. 



13 





Percentage of commercial 
shells. 


Percentage 
of culls. 


Conditions. 


Mussel beds. 


o 

a 

a 


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10 
70 
66 
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20 


11 

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45 
51 
62 


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5 


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6 
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34 

17 

37 

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84 
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92 
84 
86 


79 


.Trvnp.<? Landing 




1 
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2 
4 
2 
6 
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2 
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...do 


78 


Linton, Ky 






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...do 


75 


Donelsons Landing 






...do 


...do 


76 


Canton, Ky 


4 




.do 


...do 


74 


Eddwille Bar 




...do 


Rocks 


73 




2 


5 


...do 

. .do 




MoneyCliff 


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Mussel Shoals 




5 




3 


...do 


...do 



















Mussel Shoals was the lowest point visited on the river, but from 
reports given by the clammers the niggerhead continues to be the 
prominent shell down to the mouth of the river. 

The number of beds in this section of the river is fully equal to 
that of the preceding section, but they have not been worked as much 
because they are farther away from the center of demand and require 
transportation up the river to Clarksville. The niggerhead gains 
steadily in its percentage and at Canton passes the pigtoe, and then 
continues to increase down to the mouth of the river. There is also 
a steady decrease in the amount of culls, until at and below Canton 
nearly all the shells obtained were marketable. Of course, this 
means much to the clammer, as it does away with the necessity of 
sorting the shells and handling over the culls. 

TABULAR STATEMENT OF DISTRIBUTION OF SPECIES. 

In the table herewith given is expressed the distribution of every 
species of mussels obtained by the party in the Cumberland River 
and its tributaries. Where the mere presence of a species is all that 
is desired, it is indicated by an X. The percentages of the more 
important commercial species are indicated by numbers. The totals 
represent the actual number of specimens obtained. In order to 
catch the eye readily, all the side stations not on the main river are 
printed in italics. All commercial species are marked with an 
asterisk (*). 



14 



MUSSELS OF CUMBERLAND RIVER ANT) TRIBUTARIES. 



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MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



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16 



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MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



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20 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

RELATIVE ABUNDANCE OF DIFFERENT SPECIES. 

In forming an estimate of the relative abundance of the different 
mussels in the various beds many things have to be taken into con- 
sideration. 

For the clammer's purpose, a count of his entire catch would give 
the most reliable data, but this is usually impossible. It is almost as 
satisfactory to take the successive hauls as they come and count the 
various species in each; the greater the number of hauls counted the 
more accurate the results obtained. 

From the viewpoint of the conchologist, however, such an estimate 
is in reality only a measure of the extent to which the species in ques- 
tion is capturable by the clammer's gear, and for the following 
reasons : 

There are a number of species which never "bite" the hooks on a 
crowfoot dredge, or which do so very rarely. Such species may be 
plentiful in a mussel bed and yet never appear in the clammer's 
hauls. 

Again, some mussels are found only in small numbers and around 
the edges of a bed. The clammer makes his hauls where the shells 
are most crowded, through the center of the bed, and may miss these 
altogether. 

The clammer throws away the mussels that are too small to use as 
well as those whose shells are too thin or too highly colored. Such 
shells ought to enter into the percentages as much as the more valu- 
able species, but they do not appear in the clammer's hauls. 

Different methods of clamming produce very different results in 
the proportion of shells obtained. The crowfoot dredge, the rake, 
the tongs, and wading each secure an unduly large number of some 
species and an unduly small number of other species. 

To enumerate all the shells obtained by all the methods would give 
the most accurate results, but that is obviously impracticable. When 
the water is low the clammer gets quite a different proportion of 
species, and may even get different kinds of mussels from those 
obtained when the water is high. 

Each of these considerations has been kept in mind while making 
out the percentages; the clammer's hauls were counted; all the piles 
of culls were carefully examined; all the specimens possible were 
secured by wading along the edges of the beds; account was taken of 
the various shells found in muskrat piles; the relative stage of water 
was noted, and, so far as could be done, allowance was made for it. 
Then, too, there has been a careful consideration of numerous cir- 
cumstances which can not be shown to the reader, but which result 
from the authors' experiences at the different stations. Notwith- 
standing all these efforts, the numbers must still be regarded as 



MUSSELS OP CUMBERLAND RIVER AND TRIBUTARIES. 21 

approximate rather than absolute. But, even so, they will be of 
service to the mussel fishermen, for whom they are primarily intended. 
Only a very small percentage of the shells seen and handled could 
be kept for the final collection. 

An endeavor was made to retain typical specimens of each species 
encountered, and also all puzzling and aberrant forms, since the latter 
add much to the actual knowledge of a species, though they may 
render positive identification more difficult. 

SUMMARY OF MUSSEL DISTRIBUTION. 

The practice of the Bureau of Fisheries in examining a river and 
its tributaries from source to mouth, in regular order, throws unex- 
pected light on the distribution of species which could be obtained 
in no other way. The fauna of a river has a coherence never found 
and not to be expected in an artificial division of the country, such as 
a township, county, or State, whose boundaries are purely arbitrary. 
The larger the river and the more thoroughly the main stream and its 
tributaries are examined the more illuminating become the results. 
The study of the entire fauna of the Cumberland River and its tribu- 
taries leads to the following general conclusions, which are amply 
confirmed in all the river faunas that have been examined: 

1. When two closely related forms differ essentially in their degree 
of inflation, the flatter and less inflated one will be found in the upper 
portions of the river and in the tributaries, while the rounder and more 
inflated one is confined to the lower portions of the main river, where 
there is a weaker current and more mud. To this there are, however, 
some noteworthy exceptions, such as Symphynota complanata. 

2. The swiftness of the current, the size of the stream, and the 
kind of bottom affect other shell characters besides that of inflation. 
Consequently, where there is a mixture of conditions there is also a 
mixture of characters, and two species which in other localities may 
be well defined and easily separated will be found to merge imper- 
ceptibly' into each other. In a miscellaneous collection of shells it is 
easy to find the blue-point (Quadrula undulata) from one stream and 
the three-ridge (Q. plicata) from another, the southern mucket 
(Lampsilis ligamentina gibba) from one locality in a State and the 
pocketbook (L. ventricosa) from another. But when specimens of the 
entire fauna of a river are spread out on a table in order from the 
source to the mouth there is found such a mingling of characters that 
it is often a mere matter of individual judgment to determine some 
of the species. This is essentially true of Q. undulata and Q. perpli- 
cafa in the upper portions of the Cumberland. 

3. There is sometimes a peculiar similarity in the faunas of widely 
separated tributaries, where the conditions at first would seem to be 



22 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

very different. Such a similarity is found in Roaring and Rock 
Castle Rivers, although the localities are widely separated and the 
surrounding country quite different. 

4. Some species demand peculiar conditions, and their presence 
or abundance in any locality depends on the presence and extent of 
the favorable conditions. 

The washboard (Q. Tieros) lives in holes or depressions in the bottom, 
full of soft mud. Any mussel bed in the Cumberland that has such 
holes will be likely to contain washboards, whether that bed is high 
up the river or low down toward the mouth, and the percentage of 
the washboards will depend on the area covered with such holes. 

5. The Cumberland is very different from the Maumee and Kan- 
kakee Rivers in that it shows a marked differentiation between small 
and large stream species, between the main river and its tributaries, 
but there is very little evidence of migration along the main river 
itself. 

Such species as are confined to the upper, middle, or lower portions 
of the river owe their habitat chiefly to the fact that here, as else- 
where, they frequent smaller or larger streams, as the case may be. 

Accordingly, we may distinguish the following classes : 

(a) Small-stream species restricted to the upper portions of the 
river and its tributaries. Here belong seven species. Anodontoides 

ferussacianus was found only in the tributaries and not at all in the 
main river. The other six species, Lampsilis perdix, multiradiata, 
orbiculata, and punctata, and Alasmidonta minor and trunmta are 
distributed in various tributaries and in the main river both above 
and below the falls. None of these are commercial species. 

(b) Large-stream species, restricted to the lower portions of the 
main river. There are nine of these species, seven of which are not 
found in any of the tributaries, viz: Lampsilis ventricosa&ndfattaciosa, 
Obovaria retusa and ellipsis, and Quadrula heros, ebena, smdfragosa. 
The other two species, Lampsilis anodontoides and Quadrula undata, 
were found in Harpeth River and the former also in Red River as 
well as in the main Cumberland. The most of these large-stream 
species are good button shells, as would be expected. Indeed, the 
only exception is Obovaria retusa, which is the smallest of them all and 
for that reason the least valuable. 

(c) Species of universal distribution, which are well scattered 
throughout the entire length of the main river. There are seven of 
these species, three of which, the Ohio River pigtoe (Quadrula 
obliqua), the pink warty-back (Q. tuberculata) , and the butterfly 
(Plagiola securis), are not found in any tributary. The other four 
are the southern mucket (Lampsilis ligamentina gihba), the pocket- 
book (L. ovata), the spike (TJnio gibbosus), and the elephant-ear 



MUSSELS OF CUMBEELAND RIVER AND TRIBUTARIES. 23 

(U. crassidens) . The last two, of course, are culls, but all the others 
are valuable commercial shells. 

(d) Species confined to restricted areas, including all of the rare 
forms that are of interest chiefly to the conchologist. These include 
ail of the Truncillas, which were found in places widely separated from 
one another, and one of which was new to science; nine species of 
Lampsilis — teniata, picta, lienosa, vanuxemensis, trabalis, parva, glans, 
Jseriss-ima, and leptodon — all of which are too small or too thin-shelled 
to be of any value. Dromus caperatus and Sympliynota complanata; 
two Anodontas, imbecillis and grandis; two Pleurobemas, clava and 
crudum; and four Quadrulas, undulata, tuberosa, rubiginosa, and 
granifera. These last four have some commercial value but not very 
much. 

6. The great bulk of the mussel fauna of the river is thus made up 
of the seven universally distributed species, and two of the large 
stream mussels — Quadrula Tieros and Q. ebena. All the others are 
confined to such restricted areas or occur in such small numbers as to 
possess only an incidental or accessory value. 

NOTES ON THE VARIOUS STATIONS. 
THE UPPER RIVER AND ITS TRIBUTARIES. 

This portion of the river was examined by Mr. Boepple in 1910 as 
well as by the present party in 1911. Both the river and its tribu- 
taries are rather swift mountain streams which are much used as a 
source of power to run small gristmills, and hence they are fre- 
quently interrupted by dams. The bottom is mostly bedrock 
sandstone, with occasional fissures and sand and gravel pockets and 
bars, the latter furnishing the only localities where mussels can five. 
Consequently the shells are very few in number and widely scattered. 
The Clear Fork has more sand bars and pockets than the main river, 
and hence considerably more mussels. 

Mr. Boepple in his notes called attention to the apparent presence 
of acids in the water above the great falls, which quickly dissolved the 
nacre of dead shells, and the present party observed the same thing. 
Moreover, in the small beds above the falls the muskrats had made 
considerable inroads into the mussel fauna. Against so many 
unfavorable conditions the mussels find it very hard to hold their 
own, and the few species able to survive are not of any importance 
either to the pearlers or the button manufacturers. These mussels 
above the falls are not only thin-shelled but are much dwarfed, and 
Unio gihbosus, the most common species, has a very pale nacre, which 
frequently becomes white or yellowish and approaches closely a 
dwarfed form found in Green River, Ky. 
4788°— 14 4 



24 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

THE RIVER BELOW THE FALLS AND ITS TRIBUTARIES. 

Not only were there a great number of additional species below the 
falls, but there was also a change in the character of the shells. This 
was especially noticeable in Unio gibbosus, which was no longer a 
pale-nacred dwarf, but was of normal size and color. The mussels 
are usually found crowded about the base of the large rocks along 
the bottom of the river just below the falls. They are easily acces- 
sible to their enemies, especially during low water, and many of them 
are killed by muskrats, raccoons, mink, and occasional otter. But 
the relative number lost in this way is very small when compared 
with the corresponding loss above the falls. Hinge pearls (baroques) 
are common in this portion of the river, especially in the pocketbook 
(Lampsilis ovata), nearly every specimen of which contains a few. 
The river from Anvil Shoals, 1 mile below the falls, to Burnside was 
not investigated either by Mr. Boepple in 1910 or by the present 
party in 1911, but it was reported by a mussel fisherman to be full 
of excellent button shells. The bottom is much too stony for any 
kind of gear, however, and it would be necessary to collect the mussels 
entirely by hand. Pearling has been conducted actively along this 
portion of the river, and piles of shells left by the pearlers were 
frequent along the shore. Indeed it was reported that pearling had 
practically cleaned out the river for the first 10 miles above Burnside. 
There are two tributaries, both from the north, which enter the 
Cumberland in this space between the falls and Burnside. 

Rock Castle River is the larger of the two and is nearer Burnside. 
It was examined below the ford at Livingston, Ky., July 1. The 
shores here were high and rocky and were forested with a mixture of 
deciduous trees and hemlock. The water was clear, temperature 
81°, with a maximum depth of a foot and a half. The current was 
slow (2 miles per hour) and the bottom was very rocky and rough, 
with only a few bars and patches between the rocks filled with clay. 
The flora was remarkable and wholly unlike any that we saw else- 
where. Nuphar grew along the water's edge, Myriopliyllum verticil- 
latum, a broad-leaved Potomogeton, and a small patch of Scirpus 
americanus grew in i\e shallow water, and there was plenty of water 
willow, the whole reminding one of a bit of creek in northern Indiana 
or Illinois. The mussels were excessively abundant in the sand and 
clay patches here, and in favored localities the little Medionidus 
conradicus covered the entire bottom with the elongate slits, which 
is all of the mussel that can be seen. 

Nineteen kinds of mussels were found here, but only a very few of 
them possessed commercial value, and a few miles, farther down the 
river all the species were widely scattered. This shell bed was 
markedly unlike any of those in the main river, containing some 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 25 

species that were not found in the Cumberland at all, and others 
that were quite rare. In these respects they resemble those found 
in Roaring River in Tennessee. 

Laurel Creek, a tributary of Laurel River, was examined below 
the dam at Corbin, Ky., July 3. The shores were rocky and were 
heavily wooded with a deciduous forest, mixed with hemlock and pine, 
and still supported a remarkably rich and varied flora. The dam 
cuts off the upper portion of the river, and no mussels were found 
above it. There was a city dumping ground near at hand and the 
water was milky in color and covered with a greasy scum. Below 
the dam the bottom was very irregular and mostly solid rock, full of 
potholes and patches of sand and destitute of vegetation. 

We had expected to find a rich and varied fauna, something like 
that of the Rock Castle River, but could discover only five species, 
and three of these were represented by a single shell each. This river 
thus has almost identically the same species as the Clear Fork and the 
Cumberland above the falls. The poverty of species is doubtless due 
to the smallness of the stream and the general unsuitable conditions. 

There was no dwarfing of the species, but there were several 
peculiar modifications in the color of the nacre which were not found 
in the main river. These suggest that while there is some inter- 
course with the Cumberland there is very little interbreeding. 

The Big South Fork flows into the Cumberland at Burnside, Ky. 
Our party examined it first opposite Parkers Lake, where there is a 
fish trap and a low dam. The shores there were high limestone 
cliffs, the water was very clear, and the bottom was coarse gravel 
covered with bowlders and great angular fragments of rock, with 
some sand between them. Dead shells, recently killed by rnuskrats, 
were abundant on the rocks and on the dam at the fish trap. Twenty- 
eight species were obtained here, but although seven or eight of them 
were good button shells, they were not sufficiently abundant to make 
the gathering of them profitable. At Sloans Shoals, 6 miles from 
Burnside, during the autumn of 1910, Mr. Boepple found about 20 
species, securing them all with a rake. At the riffles, 2 miles above 
Burnside, the present party found large but rather scattered beds of 
mussels, by far the greater number of which were noncommercial. 
There were 32 species in all, and evidently some of them had yielded 
good returns in pearls, for there were many piles of shells along the 
river bank and the bed had been thoroughly worked over. 

Minute marginal cysts were abundant in the edge of the mantle of 
Vnio gibbosus, often leaving small pits along the margin of the shell. 
Baroques and the distomid of Kelly were found in Quadrula tuber- 
culata, and a few large Atax in Symphynota costata. Several of the 
U. gibbosus and two of the Pleurobema were gravid. The latter 



26 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

had fine red eggs in all four gills and the body was orange ; the former 
had coarse white glochidia in only one pair of gills. 

On proceeding down the main river from Burnside the first mussel 
bed of note is on the bar below the mouth of Fishing Creek. Very 
few living mussels were seen here, but the entire river bed was covered 
with shells which had been killed by pearlers. A large number of 
beautifully marked univalves were present among the dead mussel 
shells. 

At Fords Island the bottom of the left chute, which we examined 
most carefully, is a shingly gravel, in which it was difficult to find 
the mussels. Mr. Boepple, who examined this bed in 1910 with a 
mussel rake, reported an "almost unbelievable quantity" of TJnio 
crassidens. The present party would probably have obtained many 
more mussels if the bed could have been examined during low water. 

Four miles farther downstream, at Mill Springs, is another long 
and straggling mussel bed, which covers several miles of the river 
bottom. The latter is here composed of shingly gravel, with some 
sand bars, and is largely covered with water-willows. 

The pearlers' piles along the banks opposite this bed were chiefly 
the shells of TJnio crassidens (elephant-ear) with some Dromus and 
Quadrula obliqua (Ohio River pigtoe). Although this was not an 
important shell bed it was noteworthy for the increase in the number 
of species. The pocketbooks (L. ovata) found here were the first 
typical ones seen. 

At the pearling camp 1^ miles below Eadsville or Lock 21 we found 
the water about 2 feet above normal and rising rapidly, with a 
swift current over a gravel bottom. The pearlers were farmers from 
near by, who carried on pearling at odd times. They had thrown 
their opened shells back into the river, and there were about a ton 
and a half of them lying in the shallow water along shore. The 
pocketbooks (L. ovata), muckets, and elephant-ears were the most 
numerous species. Mr. Boepple investigated Gands Island, in this 
vicinity, and found the mussels, especially TJnio crassidens, abundant 
on both sides of the island, an unusual circumstance. 

Beaver Creek is a small tributary of the Cumberland from the 
south, opposite Rowena, Ky. This creek was investigated for a 
mile, up to a series of long riffles. The bottom was rocky with con- 
siderable mud and sand, in which were obtained a surprising variety 
of shells for so small a stream, as is shown in the table. 

In the mouth of Goose Greek, a little way down the river, a man 
was seen actively pearling with a fork. He said that he was getting 
mostly elephant-ears and that there were plenty of muckets on 
the other side of the river but the water was too high to work them. 
Mr. Boepple saw a fine lot of about 50 pearls in Rowena during his 
stop there in 1910. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 27 

Indian Creek Shoals, 53 miles below Burnside, is one of the most 
interesting mussel beds of the upper river. We found the water 
clear with a swift current over a gravelly bottom. Near the water's 
edge was a pile of about 300 pounds of shells left by a pearler. 
These were mostly pocketbooks and muckets, but contained a good 
sprinkling of sand-shells, Dromus, and monkey-faces. Mr. Boepple 
obtained a good collection of shells from this bed in 1910 and also 
from Copper Island a little farther down the river. 

Snows Island is a large island covered with coarse pebbles, upon 
which many dead shells had drifted, while others along the shore 
had been freshly killed by muskrats. At the head of Weeds Island, 
a little way below, there Avas about a ton and a half of shells left by 
pearlers, chiefly the southern mucket and elephant-ear. 

At Tear-coat Bar on July 20 the water was muddy and high from 
a heavy rain the night before. The bottom here is black gravel 
mixed with yellow sand. Out of a ton and a half of shells left 
here by pearlers about 90 per cent were southern muckets and 
elephant-ears and the remaining 10 per cent an admixture of other 
species. 

Selfs Bar contained a large and populous mussel bed which had 
been the center of active pearling operations. The 3 tons of shells 
left by them contained about the same percentage of shells as at 
Tear-coat Bar. 

Marrowbone Creek, a small tributary from the north, was examined 
up to the first riffles, a mile or more, but contained no mussels. In 
general the northern tributaries of the Cumberland were rather bar- 
ren, while those from the south were well populated. On the top 
of a hill near the mouth of this creek was an old shell pile left by 
the Indians, and from this point these shells became quite frequent, 
especially near the sites of old camping grounds. 

At Champs Shoals pearling was being actively carried on, and 
there was a large pile of discarded shells, two-thirds of which were 
elephant-ears, while nearly all of the other third were southern 
muckets. The river here widens out considerably, and there i s 
more clay and sand on the bottom. The shell bed continues with 
some interruptions from this bar down to Burkesville. At Tobins 
Landing, below Burkesville, Mr. Boepple obtained a fine collection 
of shells, representing at least 14 species. 

At Cloyds Island, below Tobins, there is an unusually good mussel 
bed which has been much worked by pearlers. The banks along 
both sides were fairly covered with the shells left by them, prin- 
cipally southern muckets and e.lephant-ears. In this bed the mus- 
sels were thickest where the current was strongest. 

Biggerstaff Bar and Island were examined July 24; at the head 
of the island were a few shells among which were found specimens 
of Lastena lata, a rare species. 



28 MUSSELS OF CUMBEELAKTD BIVEB AND TEIBUTABIES. 

A few rods below the bar there were several good-sized shell 
piles left by rnuskrats, from which we obtained an exceptionally 
fine lot of butterfly-shells (P. securis). From Martinsburg to Celina 
there were a few pearlers' piles which increased in size and number 
of shells as we approached the latter place. 

The Obey River, a tributary from the south which enters the Cum- 
berland at Celina, Tenn., and the Cumberland itself in the vicinity of 
Celina, were examined by Mr. Boepple in 1910 and again in 1911. He 
covered the lower 26 miles of the Obey River, beginning at Grass Lot 
Shoals, where no mussels were found. At Martins Bar a large collec- 
tion was obtained representing 22 species, of which the southern 
mucket and the pocketbook were the most abundant. The bottom 
here was firm coarse gravel. At Holmes Bar 24 species were secured, 
the southern mucket being still the most abundant. The current was 
swift and the coarse gravel bottom was covered with a rich vegeta- 
tion, in which the mussels were especially abundant. The southern 
mucket is the only shell in this river worthy of commercial considera- 
tion, the others being too scarce. Mr. Boepple estimated that when 
niggerheads are worth $30 per ton these muckets would be worth $50. 

From 12 to 15 years ago there was considerable pearl fishing on the 
Obey River, and a local firm said that then one could easily get a 
wagonload of mussels a day. But now the larger mussels are gone 
and the small ones have only small pearls. Fourteen of these pearls 
which were examined weighed from 2 to 4 grains each, but were of 
extra quality. 

In the Cumberland, 1 mile below Celina, there is a fair-sized mussel 
bed which has been worked for 10 years, entirely for pearls. The 
most valuable commercial species is still the southern mucket, and 
this is also regarded as the best pearl bearer. 

Mr. Boepple examined a large bed near Butlers Landing and 
secured 13 species, but the specimens were all too badly eroded and 
spotted to have any commercial value. A storekeeper here had a 
number of pearls which he had taken in trade, and he showed us an 
assortment of 4 purple, 5 yellow, and 8 white ones, of the rosebud 
type, all of which had an exceptionally good luster. 

About 3 miles below Butlers Landing we found the first pile of 
commercial shells we had seen, but they were all old shells, since no 
active clamming had been carried on for two years. There were 6 or 
7 tons in the pile, most of them of second quality, the Ohio River 
pigtoe being the most common, with the southern mucket and the 
Cumberland pigtoe (Q. cooperiana) close seconds. There were fully 
2 tons of culls, 98 per cent of which were elephant-ears and the purple 
warty-back. Mr. Boepple secured a fine collection of shells from this 
bed with the crowfoot dredge, and among them were 3 specimens of 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 29 

Lampsilis fallaciosa, the slough sand-shell, which were the first 
obtained during our survey of the river. 

At Brimstone Island there is a large mussel bed in water from 2 to 
8 feet deep, with a bottom of coarse gravel, sand, and clay. Com- 
mercial clamming had been in operation here only a few days before 
our arrival, but must have been carried on during previous years, as 
evidenced by a pile of button shells on the bank containing fully 
20 tons. 

At Carsons Bar there is another large mussel bed in water from 3 
to 6 feet deep, with a moderate current and a hard gravel bottom. 
This bed is worked only occasionally by local fishermen chiefly for 
fish bait and pearls. 

Roaring River, a tributary from the south which enters the Cum- 
berland just above Gainesboro Landing, was examined several miles 
above its mouth on July 28. Only one small mussel bed was found 
along the shore under the shade of the overhanging trees, in 3 to 6 
inches of water on a gravelly bottom. The presence of a large amount 
of Potomogeton and the abundance of Medionidus conradicus was a 
strong reminder of the Rock Castle River at Livingston, Ky. The 
abundance of Lampsilis glans was also noteworthy, since this species 
was not found anywhere in the main river. 

At Gainesboro Bar there is a small mussel bed which can not be 
worked with a crowfoot dredge, since the bottom is composed of flat 
rocks with gravel pockets in the cracks. At the lower end of the bed, 
where the rocks were well covered with a blue clay, the mussels were 
of especially fine quality, but the bed has never been fished com- 
mercially. 

We reached Salt Lick Island when the water was low and the 
mussels were moving about actively. Similar conditions were found 
at Half Pone Bar (see p. 33), and the extremely interesting collections 
obtained at each of these stations show what a remarkable difference 
a low stage of water makes in the results of collecting. There is no 
reason for supposing these two beds to be exceptionally good, and 
probably most of the beds in the Cumberland would have nearly if 
not quite equaled them if the conditions under which they were 
examined had been equally favorable. This Salt Lick Island bed 
was especially noteworthy for the large numbers of TrunciUa that 
were obtained. No parasites were found on any of the mussels. 
Lampsilis gracilis was gravid (July 31), while L. Ugamentina gibba 
and L. orbiculata approached each other so closely in all their shell 
characters as to be indistinguishable except by the color of the nacre 
and epidermis. 

At Fort Blount Bar there is a large mussel bed in water from 4 to 
6 feet deep, with a swift current over a bottom of firm gravel mixed 
with yellow clay and sand. Two men from the Ohio River had been 



30 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

working here for a week before our visit, and two more began on the 
day of our arrival. The Ohio River pigtoe is the most common but- 
ton shell. 

At Granville our party was caught in a very heavy rain, almost a 
cloudburst, and went from there down to Carthage on high and 
turbid water which rendered any satisfactory mussel survey impos- 
sible. 

Sullivans Island was investigated by Mr. Boepple when the condi- 
tions were more favorable. He found a large mussel bed in a strong 
current on a bottom of rough gravel and yellow clay. Although hq 
secured 22 species, and among them a large number of Ohio River 
pigtoes and southern muckets, the bed is worked only for fish bait 
and pearls. Two small beds at Buffalo Bar and Sand Shoals are not 
of commercial value. 

Caney Fork, one of the most important tributaries of the Cumber- 
land, joins the latter river just above Carthage. In Buffalo Valley, 
near Flat Pond, July 27, Mr. Boepple found a mussel bed covering the, 
entire width of the fork and 1£ miles long. He used a crowfoot 
dredge and scissors fork in water 5 to 10 feet deep on a bottom of 
coarse gravel mixed with sand and yellow clay. This bed has been 
fished for pearls and baroques during the last 15 years, and according 
to accounts it has yielded well. None of the shells have ever been 
sold, and fully a carload of merchantable species was seen scattered 
along the banks. 

At Rock Springs there is a much smaller bed in a swift current, 
with water 2| to 8 feet deep, the bottom being flat rocks on one side 
and much fine sand and gravel on the other. This bed has also been 
fished for 15 years for pearls and baroques, and while the shells are 
exceptionally good for button purposes they have never been utilized. 
The spectacle-case (M. monodonta) was once common here, but has 
been nearly exterminated by being used for fish bait. Another bed 
at Lancaster Island shows similar conditions; the button shells are 
of first quality, but have never been utilized. 

At the lower end of Goodall Island in the main river below Carthage 
there are two small beds separated by a short interval. The current 
is slow but steady, while the bottom is of firm gravel mixed with 
yellow clay. There was a pile of about half a ton of shells here. 
Down nearer to Lock 7 there is a third bed in water from 14 to 16 
feet deep, which was fished for pearls up to 1908, two years before the 
lock was finished. The Ohio River pigtoe is the principal commercial 
species here, with a good sprinkling of second-grade button shells. 
The effect on this bed of the dam at the lock seemed to be to kill off 
the mussels at the lower end, but to allow the upper end to broaden 
out considerably. The clammer here opened all his shells with a knife 
instead of steaming them, since he was working principally for pearls. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 31 

He was reported to have found three during the preceding week, one 
of which sold for $100. 

At Beasleys Shoals there is a large and important shell bed with 
several good-sized piles of shells along the banks. These piles aggre- 
gated about 10 tons, and the Ohio River pigtoe furnished 80 per cent 
of the merchantable shells in them. They represented chiefly the 
residue of a great amount of clamming done here in the past. An 
Ohio River clammer had taken out 200 tons of good shells and left 
about 8 tons of culls, of which the elephant-ear formed 90 per cent. 
The bottom was gravel mixed with yellow clay and covered with 
12 to 16 feet of water. Of 5 pigtoes examined 4 were gravid, 2 had 
young in the outer gills only, while the other 2 had a number of young 
in the inner gills also. The Quadrula subrotunda had orange flesh 
while part of the gills contained carmine eggs, most of which had 
been aborted. 

Below Cedar Bluffs we found a pile of 12 tons of shells which had 
been collected a year or more before, and cribbed. The mussel bed 
here was large with a very slow current over a bottom of gravel 
covered in some places with clay. The bed has been extensively 
fished for pearls; during the previous year (1910) 8 boats had been 
employed and they collected over 100 tons of shells, more than half 
of which were saved and sold. But there was fully a carload of good 
button shells scattered along the banks. 

Goose Creek, a tributary of the Cumberland from the north, was 
examined August 10, but although the conditions seemed in every 
way favorable no mussels could be found. 

At Daniels Landing the mussel bed is half a mile long and 150 feet 
wide in water 12 to 16 feet deep, with a bottom of yellow clay and 
sand changing to rocks at the lower end. The fishing here has been 
chiefly commercial since pearls are scarce. Eight men fished this 
bed in the summer of 1910 and obtained 100 tons of shells, the prin- 
cipal commercial mussel being the Ohio River pigtoe, which is of 
extra-large size and of the best quality., A few very large nigger- 
heads were also found. In spite of the large amount of shells taken 
from this bed it still remains one of the richest in the river. 

At the mouth of Spring Creek, below Hunters Point, there is a 
large mussel bed 1 mile long and 125 feet wide, in a very slow current 
over a bottom of gravel and yellow clay covered in places with mud. 
This was first fished in 1910, when 50 tons were taken; at the time 
of our visit in 1911 the clammers had obtained about 14 tons, nearly 
all of Ohio River pigtoe, with a few washboards and niggerheads. 
Another large mussel bed was reported at the foot of Wings Eddy 
Bar, and still another at Armstrongs Island. At Cairo we saw a pile 
of 12 tons of shells, mostly Ohio River pigtoes. 
4788°— 14 5 



32 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

At Grallatin Landing the mussel bed is lh miles long and from 40 
to 60 feet wide, on a bottom of gravel and yellow clay covered with 
mud. The river widens considerably, there is much dead water, and 
the shores are low, making the conditions almost lake-like. This is 
all the result of excessive backwater from the lock dam just below. 
The first Quadrula fragosa was found here. 

At the head of Lindsleys Island we found a very large number of 
small shells killed by muskrats; 95 per cent of these shells were pig- 
toes. There is no commercial fishing here nor even any pearling. 
We found in this bed our first yellow sand-shell, and also a spectacle- 
case, specimens of which we had not seen for some time. Farther 
down the river, at the end of Lindsleys Bar, there was a clammer's 
camp. About 600 pounds of shells had been collected, of which the 
pigtoe formed 50 per cent, the washboard 25 per cent, and the 
remainder mixed species, including a few yellow sand-shells. There 
was a good mussel bed at Hills Island above Nashville on a muddy 
bottom in a fairly rapid current. Many mussels had been killed by 
muskrats who seemed to have a particular liking for small pigtoes. 

Stones River, an important tributary from the South, was exam- 
ined along its East Fork at Walterhill, Tenn. The water was shallow 
and turbid with numerous riffles; the bottom was composed of loose 
rocks with intervening gravel bars, covered with plenty of water 
willow. 

Below the ford was found a large number (70) of a beautiful new 
species of Truncilla (see p. 46), many individuals of which had been 
killed by muskrats. The Symphynota costata found here were remark- 
ably large, and contained many lusterless pearls. 

The West Fork of this river was visited at Murfreesboro, Tenn. It 
is somewhat larger than the East Fork and is broken up by divers 
islands covered with water willows. There were many Anodonta 
grandis and Symphynota costata of large size on the bank, recently 
killed by pearlers. 

The musseWauna here is remarkable in containing several species 
not found at all in the Cumberland, and in a peculiar interchange of 
species. L. ovata of the Cumberland is replaced here by the genuine 
L. ventricosa and Q. perplicata is replaced by Q. undulata. The pres- 
ence of Q. rubiginosa is unexpected, and that of the genus Anodonta 
is interesting, since this is the only place in the Cumberland or its 
tributaries where representatives of this genus were found. 

At the foot of Gowers Island, 25 miles below Nashville on the main 
river, there is one of the most important mussel beds in the entire 
Cumberland. And we found here the largest pile of mussel shells yet 
seen, about 80 tons with 8 tons of culls. The bed is 3 miles long and 
from 60 to 175 feet wide in a strong current on a bottom of gravel 
mixed with sand and clay. The young pigtoes here were all so 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 33 

brightly rayed that for a time they were regarded by the clammers 
as possibly a new species. Harpeth River, a tributary from the 
south which enters the Cumberland a little way above Lock A, was 
examined 5 miles above its mouth. The bottom here was of shingly 
gravel, changing to solid rock and farther up to beds of soft mud. 
There was formerly a large mussel bed here, but the backwater from 
the lock dam has killed the mussels in the lower portion of the bed. 
Another large bed was reported 1| miles farther up the river. 

The unusual size and thickness of the shells obtained here suggest 
that this river would yield exceptionally good button material. The 
margins of the shells were much pitted, indicating parasites in unusual 
abundance. The presence of fine large L. ventricosa and S. costata 
so near the mouth of the river is remarkable, since both of these 
species are absent from the Cumberland. 

Below Lock A we saw numerous sites of old shell piles where clam- 
ming operations had been carried on in the past. At Half Pone Bar 
the current was swift, the water shallow and somewhat turbid, and 
the bottom firm gravel and sand. The large number of specimens 
and species is at least partly due to the peculiar configuration of the 
bottom and the low stage of the water, the conditions being similar 
to those at Salt Lick Island (see p. 29). The great majority of the 
shells obtained were young, but many of them were eroded at the 
umbones. P. donaciformis was exceptional in being very thin and 
having a pink nacre. The large number of Plagiola is noteworthy, 
together with the only specimen of Truncilla Jlorentina found below 
Nashville. 

At the Seven Mile Ferry above Clarksville the current was rather 
feeble, the water clear, and from 5 to 8 feet deep, and the bottom 
composed of fine gravel. From this point on down the river a crow- 
foot bar was employed, similar to that used by commercial clammers 
but shorter and smaller, and furnished with 50 hooks. The latter 
were of two kinds, the ordinary form used by clammers and an 
improved form invented by Mr. Boepple, having a knob at the tips 
to prevent small mussels from taking hold or larger ones from dropping 
off. Hauls were made 200 feet long, the first as near the shore as 
possible, and each succeeding one 10 feet farther out. The detailed 
record of the different hauls made at a few stations is given in full, 
in order to convey a more accurate idea of the number and distribu- 
tion of the mussels, and the ease or difficulty with which they could be 
caught. Such a record was kept for all the stations in this portion 
of the river, and forms an important factor in determining the relative 
abundance of the mussels. 

At Owl Hollow Bar, 2^ miles above Clarksville, we found a swift 
current with clear water over a clay bottom, more or less mixed with 
gravel. This bed had been worked for eight years and showed signs 



34 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



of depletion. The detailed reeord of the 14 hauls made here is given 
in the following table : 

Hauls Made at Owl Hollow Bar. 





1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 


14 






tal. 




4 


4 


4 


3 


3 


5 


4 


3 


3 
1 


6 


4 


9 


4 


6 




1 


















1 












1 
















1 














1 




















1 


1 








2 




1 


3 




1 




3 
1 
1 

7 


1 


1 




1 
1 


1 
1 




12 




1 






4 




2 
13 

2 


"6" 
i 


1 
14 
1 
1 






4 

3 


2 
.... 






fi 




6 
1 


4 


2 


3 
2 


3 

i" 


16 
3 


5 
1 

1 


4 
1 
1 


90 




17 






1 
1 


4 
















1 








?, 














1 














1 
















1 


2 


5 
1 


5 

1 


1 


1 


1 


16 


















?, 




18 


13 


17 


8 


4 


14 


10 


6 












Total 


7 


12 


11 


22 


10 


7 


159 







This was one of the most important mussel beds visited, since 
clamming was going on actively at the time of our visit, the shells 
being used at the Clarksville blank factory. The bed has been 
worked for 10 years with from three to six boats every summer, but 
it shows very little sign of depletion. In sorting the shells the wash- 
board (Q. lieros) is piled by itself, because it is badly stained, and 
sold at one-half or one-third the regular price. It forms about one- 
fifth of the entire catch. 

Of the first-grade shells the pigtoes are much the most abundant, 
followed by the niggerhead and the monkey-face. Mussel enemies 
are scarce, most of the mink and muskrats having been trapped. 
Pearls and baroques are rare, slugs run about three-quarters of an 
ounce to the ton. A large number of the pigtoes obtained were 
gravid and several had young in all four of the gills. 

At Clarksville June 12 the river was very low and a large sand bar 
was being uncovered. The bottom was fine gravel and the water 
rather shallow, with a slow current. The yellow sand-shells were 
traveling rapidly into deeper water. Plagiola donaciformis was 
gravid. 

At Red Rock Bar, below Clarksville, on June 6 the water was 
unusually clear, about 8 feet deep, and there was practically no 
current, the bottom firm gravel. Fourteen hauls were made here 
under the same conditions as at Owl Hollow Bar, save that each was 
300 feet long. The mussels found gravid here were 1 0. refiexa, 
2 Z7. gibbosus, 3 Q. perplicata, 1 Q. pustulosa, 77 Q. obliqua, and 10 
Q. ebena. Tins is the only place in the main Cumberland that we 
found S. complanata. This bed has been worked eight years and 
begins to show the effects of it. The shells obtained are of better 
quality than when the work first began, but there are fewer slugs, 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



35 



the shells being younger. Quadrula perplicata, called locally the 
"round-lake," is the pearl bearer here. 



Hauls Made at Red Rock Bar. 





1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 


14 


To- 






7 


6 


2 


6 


8 


2 


9 


12 


5 


9 


12 


14 


11 


4 


tal. 






























1 






1 






1 
1 










1 




1 










3 
























1 
















1 
1 














1 


















1 
2 
1 
3 




1 








•^ 
















1 


1 
2 










4 










1 








2 




2 

2 


8 




3 


3 




1 


2 


.... 


1 

2 




1". 




1 


2 


i 


1 




s 




1 

3 

21 

3 


1 
1 
2 
2 














2 












2 

25 
2 


'l2' 


2 
3 


1 

13 


8 
1 
1 


1 

18 


2 

10 
3 
1 


'26" 
5 


1? 




6 


17 
1 


20 
1 
1 


12 


1x7 




IS 




1 






4 




1 
3 

1 

36 






1 










2 




























3 




1 






1 


2 


5 




2 


1 










13 


















Total 


12 


7 


22 


24 


16 


37 


18 


15 


19 


11 


23 


16 


29 


o r=; 







Trices Landing is 1^ miles below Clarksville and the conditions 
are almost exactly the same as at Red Rock bar, except that the 
bed is full of "hang-ups," and therefore not fished commercially. 

At Meeks Spring bar, about 8 miles below Clarksville, some fine 
springs enter the river, one of which has its outlet richly incrusted 
with diatomaceous scum. The current was very slow and the water 
unusually clear over a bottom of coarse gravel. This bed has been 
fished for 10 years and 500 or 600 tons of shells have been taken from 
it. Most of the O. rejlexa and Q. fragosa were found cleaned at 
muskrat holes and were practically the only shells there. The yellow 
sand-shell and the rabbit's foot had been going shoreward during a 
previous rise in the river, but turned and went back when the water 
fell. Many of these sanc^-shells were gravid June 10 and were used 
in making a plant of mussels in the river at Clarksville. 

The Red River is the only tributary of any size that enters the 
lower Cumberland from the north. No mussels could be found for 
several miles above its mouth, probably because the bottom was 
found to be covered with soft mud which shifted considerably during 
high water. 

At Ringgold, on the west fork of the river, there is a high milldam, 
which backs the water up for several miles. No mussels were found 
above this dam, and below it they were rather scarce and all of small 
species. Several L. multiradiata were found which showed no rays, 
a few L. vanuxemensis, and one live L. glans. This proved to be the 
only place where vanuxemensis occurred. 

Mr. Boepple visited Port Royal, at the junction of the two forks of 
Red River, on June 14. The river here is not large and is shallow 



36 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES^ 

except a few deep holes; the bottom is gravel and mud. The mus- 
sels were collected with a rake and by wading, and were mostly near 
the bank in the mud, only a few being found in the gravel. Sixteen 
species were obtained in all, two of which, S. costata and S. compla- 
nata, were rare in the main river. The mussels were said to have 
been formerly abundant, but they had been nearly cleared out by 
pearlers, and not enough marketable species were left to make fishing 
profitable. 

Haynes Lake lies several miles below Clarksville, on the north side 
of the river, and is apparently a part of the old river channel. It is 
about a mile long and surrounded by woods; the bottom is soft mud 
and the water is about 3 feet deep, with a temperature of 89°. Very 
large specimens of Anodonta grandis gigantea were obtained, 2 of 
which contained sporocysts of some distomid, while 2 others were 
gravid (Sept. 4). The nacre of 8 was purplish, that of the remain- 
ing 17 a beautiful creamy white. Of the 2 specimens of A. imbeciUis 
1 was gravid. 

Elk Creek Shoals, 13 miles above Dover, had a current of 3 miles 
an hour in water 10 feet deep over a bottom of gravel mixed with 
some sand. Nine of the pigtoes obtained here were gravid (May 30), 
and on the land bar above the shoals was found one dead TrunciUa 
sulcata, a species which is exceedingly rare in the Cumberland. 

Walter's shelling camp was about a mile below these shoals, and 
Mr. Walter very kindly conveyed us up and down the river in his 
launch, giving much valuable information. He had a pile of shells 
containing about 150 tons, of which the most important button shells, 
in the order of their abundance, were the Ohio River pigtoe, the 
Cumberland pigtoe, the monkey-face, the yellow sand-shell, the but- 
terfly, the niggerhead, and the southern mucket. 

At Glasgow Landing, 2 miles above Dover, on May 29 the current 
was about 4 miles an hour, the water high and muddy but rapidly 
falling, and the bottom gravel mixed with clay. About one-third of 
the pigtoes were gravid, the glochidia being usually in the lower half 
of the outer gills. The niggerheads were also in the early stages of 
gravidity, all four gills being red and padlike; one elephant-ear was 
gravid. At the foot of Dover Island the conditions are the same as 
just recorded except that the water was 20 feet in depth. A small 
species of Atax, with broad white marks on the back, was found on 
several of the mussels obtained here. Marginal distomid cysts were 
fairly common, especially in P. securis (the butterfly). This same 
butterfly was frequently gravid, the pigtoe was less often gravid, and 
a single specimen of Q. fragosa had glochidia in all four gills. 

A noteworthy feature of the lower river, somewhat marked at 
Clarksville, but decidedly more so at Dover and below, is the land- 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



37 



slips that occur along the banks, when great masses of earth slide 
into the water, sometimes carrying trees with them. 

At Jones Landing there was another clammers' camp, operated by 
a Mr. Scarborough, who rendered us considerable assistance. The 
water here was 15 feet deep and the current about 3 miles an hour 
over a bottom of mud and gravel. Sixteen hauls were made here, 
with the following results : 

Hauls Made at Jones Landing. 



Number of hauls 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 


12 


13 


14 


15 


16a 


To- 


Duration in minutes. . . 


15 


8 


6 


5 


4 


2 


2 


4 


8 


6 


6 


8 


5 


8 


4 


4 


taL 
















1 


2 


















1 
























1 








1 














1 




















1 




1 


1 
1 


1 




1 
2 














1 
1 
1 


2 






7 
















1 






5 




























1 























1 










1 












1 
2 
31 
4 
1 








1 
1 

18 
.... 


1 
3 

24 
2 












T 




2 

13 

1 

1 


2 
15 


5 

12 

1 


1 
12 

1 






3 

20 
3 


"i 


6 

32 

4 

1 


5 
1 


1 

27 






?« 




14 




25 
2 




?fi8 




n 








5 
















1 










1 














1 














1 


2 
2 


.... 


4 




1 


1 
2 






1 
1 






2 






1 
1 




8 






2 


1 
1 










7 
























1 




















1 














1 
































Total 


19 


22 


19 


16 i 44 


18 1 


28 1 23 1 32 


5 


46 1 31 


31 


31 





366 

























a No mussels taken. 

At Three Sisters Springs, near Linton, Ky., some remarkably 
large springs flow out of a cave into the river. There was a current 
of 4 miles an hour in water 20 feet deep over a bottom of soft gravel. 
No parasites were found except distomid cysts along the margin 
of the mantle of a few shells. Stained and rough t*ps, which in 
some places indicate pearl formation, were common in the shells 
here. Six of the pigtoes had the lower half of the outer gills filled 
with glochidia (May 24). All the mussels examined had their intes- 
tines filled with greenish mud and appeared well fed. 

The main bed is a little below the springs and had been worked 
for four seasons. Our helper, who had been a professional clammer, 
had on one occasion dug in this bed 13 boxes of shells of 100 pounds 
per box in one day. This was in competition with another man 
who dug 12 boxes in the same time — a ton and a quarter by the two 
men in a single day. 

Below Linton shell beds are common but none were being worked 
above the mouth of Donelsons Creek. The largest of these beds is 
at Dead Mans Bar, where there was a large pile of culls near the 
mouth of Terrapin Creek. 

At Donelsons Creek a clammer had just begun working and had 
only a few shells, chiefly pigtoes, washboards, niggerheads, and 



38 



MUSSELS OP CTJMBEBLAND EIVEB AND TBIBUTABIES. 



monkey-faces. A mile below Canton, Ky., there is another bed in 
12 to 15 feet of water which had been worked previously as was 
evidenced by an old shell pile, in which a single valve of L. falla- 
eicsa was found. In the hauls here taken by our party were obtained, 
May 23, four gravid niggerheads and five pigtoes. 

The bed at Eddyville, Ky., examined May 18, was on a gravel 
bottom covered with 15 feet of water, with a current of about 2 
miles an hour. This bed had been worked more or less for four 
years, but was difficult and unsatisfactory on account of numerous 
"hang-ups." 

Just above the Ferry at Kuttawa, Ky., there was a large mussel 
bed on a bottom of sand and gravel, covered with 8 or 10 feet of 
water, with a swift current. Eighteen hauls were made with the 

following results: 

Hauls at Kuttawa, Ky. 



Number of 
hauls 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


11 a 


12 6 


13 o 14c 


15 


16 


17 


18 




Duration, 
minutes 


3 


3 


6 


4 


4 


4 


4 


3 


4 


6 


5 


2 


4 


5 


5 


2 


5 


5 


To- 
tal. 


Length in feet 


100 


200 


100 


100 


100 


30 


100 


50 


50 


100 


100 


30 


60 


40 


100 


30 


100 


100 










2 
































2 










1 










1 


















2 














1 






















1 




















1 


















1 


































1 




1 
















1 




















1 






6 
8 
2 


5 
4 


3 
3 
7 
1 
1 
1 


6 
18 
11 


1 


7 
1 
3 








1 








4 


"9' 
7 
1 


1 

12 

8 

1 


34 




1 




26 
2 
1 
2 








6 


88 






8 










48 


















4 










2 


1 


3 


1 
















10 


pustulosa 
perplicata 




















1 

8 


.... 


?, 










1 






4 
















IS 














1 
















1 


tritogonia 














1 

3 






















1 






1 


4 


1 




1 




1 














2 


3 


16 






1 














1 






































Total 


1 


17 


12 


20 


39 


3 


17 


14 


4 


34 





1 








6 4 


29 


25 


?,?6 











o No mussels, due to shifting sand. & Water 50 feet deep. « No mussels. 

Of the gravid mussels obtained in these hauls the elephant-ear ( U. 
crassidens) had the entire outer gills padlike, striate, and white. 
Lampsilis orliculata has a marsupium that is black-edged, while the 
mantle is striated brown and black like that of L. ventricosa. The 
pigtoes (Q. obliqua) were just beginning to become gravid (May 13), 
with minute white spawn along the crenate edge of the outer gills. 
In Lampsilis gracilis the posterior half of the outer gills had much 
the appearance of a lima bean, in which the conglutinates were some- 
what separated, with no black edge and no furrows. 

CHARACTER OF WATER OF THE CUMBERLAND RIVER. 

In the coal regions of the upper Cumberland River the water is 
generally clear and of an acid nature. The acidity is well shown by 
the limy parts of the dead shells being greatly dissolved away and in 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 



39 



many cases the epidermis alone left. That the mussels do not thrive 
well in this portion of the river is probably due to the fact that the 
bottom is rocky, food scanty, and the water deficient in lime. 

Below the Cumberland Falls in the limestone formations the water 
contains a considerable percentage of lime. Here the shells are much 
larger and thicker than those above the falls. 

The table given below is taken from the United States Geological 
Survey "Water-Supply Paper 236/' by R. B. Dole, and shows the 
mineral conditions of the Cumberland River, at Nashville, Tenn., and 
Kuttawa, Ky., two widely separated localities of the lower river. A 
sample of water was taken daily, these mixed, and a sample from the 
mixture was taken for analysis. There were about 3 analyses made 
per month, or 36 per year. This method gives a much better general 
knowledge of the conditions than a sirgle sample would do. From 
Nashville the samples were collected from October 24, 1906, to 
November 3, 1907, and 35 analyses made; from Kuttawa, from Jan- 
uary 11, 1907, to January 11, 1908, 34 analyses were made. 

The following table gives the general average of the analysis, in 
parts per million, and also the per cent of the anhydrous residue: 

Mineral Analyses of Water from Cumberland River. 
[Parts per million, unless otherwise stated.] 



Near Nashville, 
Tenn. 



Mean. 



Anhy- 
drous 
residue. 



Near Kuttawa, 
Ky. 



Mean. 



Anhy- 
drous 
residue. 



Turbidity 

Suspended matter 

Coefficient of fineness 

Silica (Si0 2 ; 

Iron (Fe) a 

Calcium (Ca) 

Magnesium (Mg) 

Sodium and potassium (Nan 

Carbonate radicle (C0 3 ) 

Bicarbonate radicle (HC0 3 ). 

Sulphate radicle (S0 4 ) 

Nitrate radicle (N0 3 ) 

Chlorine (CI) 

Total dissolved solids 



K). 



126 
94 

.74 
20 

.42 
26 
3.6 
9.6 

92 
14 
1.2 
2.1 
119 



Per cent. 



16.4 
i.5 

21.3 
2.9 
7.8 

37 



11.4 
1.0 
1.7 



176 
165 
.92 
18 

.30 
28 
4.3 
7.8 
.9 
100 
9.7 
1.8 
3.0 
124 



Per cent. 



14.6 
1.4 

22.8 
3.5 
6.3 

40.6 



7.9 
1.5 

2.4. 



a Fe20 3 . 

COMMERCIAL VALUE OF THE MUSSELS. 

Taking into consideration both the relative abundance of the species 
and the intrinsic value of the shell, the southern mucket (L. ligamen- 
tina gibba) is the most important commercial mussel of the upper 
river; that is, from Burnside down nearly to Nashville. 



40 



MUSSELS OF CUMBEELAND EIVEE AND TEIEUTABLES. 



From Nashville to Clarksville the mucket is not relatively as 
abundant, and is consequently surpassed in value by the Ohio River 
pigtoe (Q. obliqua). 

From Clarksville to the mouth of the river the honors are divided 
between the pigtoe and the niggerhead (Q. ebena). There are other 
shells all alorg the river which possess a high intrinsic value but are 
not found in sufficient quantities to equal the ones just mentioned. 
The most important of these are the yellow sand-shell (L. anodon~ 
toides), the most valuable of all our fresh-water species, the butterfly 
(P. securis), Lampsilis orbiculata, a shell of very high value and 
desirable for propagation, and the Missouri niggerhead (0. ellipsis). 
The Cumberland pigtoe (Q. cooperiana) and the long niggerhead (Q. 
subrotunda) are also much esteemed by the button manufacturers. 
Samples of shells from the upper portions of the river were carefully 
weighed, measured, and appraised by Mr. Boepple, with the results 
indicated in the following table: 

Commercial Value op Mussel Shells Taken from the Cumberland River in 
October and November, 1910. 



Species. 


Locality. 


Weight. 


Num- 
ber 

shells. 


Num- 
ber 
blanks. 


Lines. 


Num- 
ber 
gross 
per 
ton. 


Value 
per 

gross. 


Value 
per 
ton. 


Lampsilis liganien- 
tina gibba. 
Do 


Martinsburg, Ky 

Martinsburg, Ky 

do 


Pounds. 
5 

2J 

5 
5 
5 

} ■ 

5 

24 
2J 
2 

2J 


16 

12 

17 
17 
32 

12 

37 

ui 

n§ 

34" 
13§ 


287 

180 

222 

222 

390 

f 57 

\ 129 

76 

23 

T67 

125 

137 


20 
20 

20 

36 
20 
30 
35 

20 


685 

916 
522 
723 
928 
135 
307 
180 
109 
319 
744 

652 


Cents. 
5 

6 
3 
5 
5 

20 
7 

10 
15 
2 
2 


$34.25 
54.96 


Do 


15.66 


Do 


36.15 


Do 


Burnside to Burksville. . 
/Obey River, Celina, 
\ Tenn. 

Coes Landing, Ky 

Martinsburg, Ky 

do 


46.40 


Do 


27.00 


Quadrula obliqua . . 
Dromus dromas 


21.49 
18.00 
16.35 
6 38 




...do 


14 88 


and small Quad- 
rulas mixed. 6 


do 













o Tips. 



b Pearly tips. 



A good idea of the extent of clamming operations on the river 
below Nashville may be obtained from the following data, contributed 
by various shell buyers at Paducah, Ky. : On some of the beds 
mussel fishing has been conducted for at least 10 years. One mussel 
firm, with headquarters at Paducah, had 300 boats operating from 
Paducah to Nashville. In 1907 this company obtained 1,783 tons 
of shells from this part of the Cumberland River; in 1908, 1,400 tons; 
in 1909, 1,100 tons; in 1910, 1,125 tons. In consequence of a sudden 
drop in the price of shells this company was not working the river 
during 1911. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 41 

Another buyer reported 500 tons obtained from the same region 
of the Cumberland by his company during each of the years 1907, 
1908, and 1909, but only 100 tons in 1910. 

In addition to these companies there were Ohio River parties and 
private fishermen operating in the river, which must have increased 
the annual output to considerably over 2,000 tons per year. 

Because of the drop in prices mentioned above, none of the larger 
companies were operating the river during 1911 with the exception 
of Mr. Walter, at Dover, Tenn., and the blank factory at Clarksville, 
Tenn. 

BREEDING SEASON OF THE CUMBERLAND MUSSELS. 

Throughout the progress of the survey the various species of mussels 
were examined as to breeding condition and the date at which the 
various species were found gravid is shown in the table following. 
In addition to the table, which gives only the bare facts, the follow- 
ing additional notes will prove of interest and value. 

The only Lampsilis ovata found gravid was on May 13. Mr. Boep- 
ple sent in some gravid examples during the late autumn of 1910. 
Without doubt this species is usually gravid from autumn until the 
next spring. L. multiradiata was found becoming gravid July 28. 
In other streams we have found it fully gravid in September and 
October. Lampsilis anodontoides was found fully ripe in abundance 
from June 10 to 21. The breeding season of this species is well 
known; it usually becomes gravid in autumn and remains so during 
the winter. Quadrula perplicata was noticed becoming gravid May 
24, and gravid samples were still found July 27. Although Quad- 
rula cooperiana remained gravid for a considerable length of time, 
we saw only a few samples; the citations refer to single individuals, 
so that, while we have it recorded from June 3 to August 11, 
only 11 gravid examples altogether were seen. The characteristics 
of the gravid mussel are described under the discussion of the species. 
It is a desirable species to propagate. Quadrula obliqua is the most 
prolific mussel in the river, and we saw many more gravid examples 
of this than of any other species. From June 3 to 10 is the height of 
its breeding season, and at that time about half the catch obtained 
would be gravid. When the life history of the species is known and 
the fish which serves as host, it will be easy to procure material for 
propagation during a considerable part of the summer. 

Quadrula ebena was observed in early stages of gravidity about the 
beginning of the work, and gravid examples were obtained as late as 
July 16. The other species noted are not of special economic impor- 
tance and gravid examples were found only in small numbers. Suffi- 
cient information about them can be obtained by a glance at the 
table. 



42 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

Table op Gravid Specimens op Mussels Found in the Cumberland, 1911. 



Dates. 


> 
o 


03 

.a 

1 


5 
'3 

3 
O 
— 

o 

a 

03 

-i 


.2 

a 

= 

-3 


■1 

o 

X 


B 

i 



o 


I 

.3 

o 

d 


■1 
6 


53 

d 


i 

a 

o 

1 


-3 


1 

o 

— ' 

d 


3 

53 

o 

d 
X 


53 

o 

3^ 


53 
03 

a 

& 


03 

1 

o 

d 


53 

l> 

» 

PI 

S3 
(S 

a 

d 


03 

o 

5 
B 

d 


o 
tffl 
53 

d 


a 

.53 

3 
- 
O 

o 

o 

d 


53 

3 

o 

d 

X 
X 


si 
o 

.3 
o 

o 
o 

d 


03 

"3 

o 

M 

& 
g 

d 


I 

d 


i 

S 

s 

d 


Mav 13 


X 














































X 










































X 






18 


































X 
















23 
















X 






























X 
X 




24 
















X 












X 












X 

y 








29 




































x 




























X 
X 








X 




X 




X 
X 


X 


X * 






X 
X 


V 


5 


















.., 




X 


X 


X 
X 
X 
X 




6 . 
































7 
















































10 .. 






X 












































16 .. 
















X 
































17... 






X 
























X 




















21... 






X 










































28 






















X 
X 
X 




























29 


















































Julv 1 












X 














X 


X 






















8 






























X 
X 


X 
X 

X 
X 

X 

X 
X 
X 
X 










9 














X 
















X 
X 










X 








10 










































11 






































X 
X 
X 






X 




13 
























X 


















14 






































y 




X 
X 




15 










































16 








































X 




27 






























X 


X 












X 




28 




X 




































31 








X 










































Aug. 9. . . ; 








































X 
X 
X 
X 

X 
X 










10 








































X 
X 

X 




X 






11 










































17 
















































IS 
















































21 


















































Sept. 2 


















































3 




















































4 




















































9 




















X v 


























































! 





















PEARLS AND PEARLING IN THE CUMBERLAND. 

Just when pearling began on the Cumberland there is no definite 
record. It has been in operation quite steadily on the upper river 
for at least 20 years. It is not generally carried on actively the year 
round, but chiefly in August and September, when the water is low. 
There are few professional pearlers, however; that is, men who devote 
their entire time to the gathering of pearls. Most of the pearling is 
carried on by farmers at odd times, and by men who in the winter 
devote then energies to lumbering, chopping, or trapping. 

Hunting for pearls is confined mostly to the upper river and the 
tributaries. It seems that the conditions suitable for pearl formation 
are more abundantly fulfilled in small streams. 

The first sign of active pearling operations seen by the present 
survey was encountered about Burnside. The search for pearls 
extended above the town as far as Seven Mile Shoals and downstream 
as far as Celina, and less actively to Carthage and beyond. A short 



MUSSELS OF CUMBERLAND KIVER AND TRIBUTARIES. 43 

distance below Burnside pearling has been recently in active opera- 
tion, at Pittman and Fishing Creek. From Burnside down to 
Burkes ville Mr. Boepple had noted in 1910 that the river bed was 
well filled with shells killed by pearlers, and in 1911 the same work 
was being continued farther on downstream. At Patty Shoals below 
Mill Springs in 1910 "yellow mussels" (L. ligamentina gibba) had been 
pretty well fished out, since the pearlers opened only this species. 

In order that due allowance may be made for the inevitable degree 
of unfounded rumor on such subjects, we will give at first the reports 
of the rivermen and supplement them later by our own observations. 

At Burnside we heard that a pearl had been found at the mouth of 
Pittman Creek which was worth $250 or $300 and another that had 
been sold for $40, and we were told of a man living down the river, 
back some distance in the country, who had a fad for pearling and 
buying pearls, and who had accumulated in this way about $20,000 
worth of pearls, baroques, and slugs at the time our informant 
visited him. Many pearls had been found in the vicinity of Eadsville, 
the highest price any single pearl from that locality brought 
being $800. 

In August and September 100 men were often pearling at once on a 
shoal near Rowena, and the highest price paid for any single pearl 
was $500. There had also been much pearling on a mussel bed below 
Tear-coat Bar and on another at Clouds Island during the past five 
years, sometimes as many as 50 men working at the same time. At 
Goodall Island, for 20 years previous to the time of closing the 
lock, pearling had been in active operation. At one time 150 men 
were at work together on the bed, and in one week $30,000 worth 
of pearls were found. Pearling had also been carried on near the 
mouth of Goose Creek above Hartsville in former years, but it stopped 
after the building of the lock below, which flooded the beds with lock 
water and rendered it difficult to obtain the mussels. 

Not only the upper river but its tributaries also were famous for 
pearls. At Carthage it was said that better pearls were found in the 
Caney Fork than in the Cumberland, and that they commanded a 
much better price. Mr. Boepple, who investigated the lower 26 
miles of the Obey River at Celina, remarks: "Twelve to fifteen years 
ago there was much pearl fishing here, and it seems to have paid 
until, indeed, the mussels had been fished out by pearlers." Stones 
River was in good repute as a pearling stream, and a merchant at 
Clarksville stated that his father used to buy many pearls from there. 
There had been active pearling on this river only a short time before 
our visit, and some shells left by the pearlers and examined by our party 
showed indications of pearl formation. Red River, which enters at 
Clarksville, is said to be a good pearl-bearing stream in its upper 



44 MUSSELS OE CUMBERLAND RIVER AND TRIBUTARIES. 

portion, and we saw a number of very good pearls from there. Little 
River, across from Canton, Ky., is also said to yield numerous pearls, 
which, however, are rather small. 

Our own observations, as well as the records of people engaged 
in the pearl trade, indicate that pearling was once an important 
occupation in the upper river. We saw in many places large piles 
of shells left by pearlers along the river banks, and came across one 
party actively engaged in pearling. Mr. Boepple saw a collection 
of pearls in Rowena valued at $1,000, and this represented only a 
portion of those found in the vicinity, since the largest and finest 
pearls were sent directly to New York. At Butlers Landing a store- 
keeper showed us a very pretty collection of "rosebud" pearls, all 
with a good luster, four of which were purple, five yellowish, and eight 
white. 

At Clarksville, as mentioned above, we saw some very pretty 
pearls from Red River. One of the principal merchants at Car- 
thage buys about $15,000 worth of pearls every year. The highest 
price he had paid for a single pearl was $2,500. They generally 
range from $20 to $300. A shell buyer at Paducah, Ky., bought 
$2,000 worth of pearls during the season of 1910. While genuine 
round pearls are not common in the lower river, rough pearls and 
baroques are usually present to the amount of three-fourths ounce 
per ton of shells. The baroques vary from $2.50 to $3.50 an ounce. 

On account of the ground to be covered and the time at our dis- 
posal, together with unfavorable weather while on the upper Cum- 
berland, we did not have opportunity to devote very much atten- 
tion here to pearl formation, though this region would prove an 
exceptionally good location from which to attack the problem. In 
looking over the flesh of some mussels recently killed by pearlers a 
number of black distomid cysts, similar to those found in the Maumee 
River (Indiana and Ohio), were observed, and these probably figured 
in part at least as an exciting cause. 

A study leading to the discovery and the consequent conserva- 
tion of the peculiar conditions which favor pearl formation in the 
upper Cumberland and its tributaries, accompanied with active 
propagation of the southern mucket in that region, would be highly 
desirable. The mussels are not yet so nearly exterminated that 
gravid material can not be readily procured, though it is feared 
that they soon will be, in view of the active depredations of the pearl- 
ers. It is believed that mussel planting could be kept well ahead 
of any onslaught likely to occur and that the resulting harvest 
would yield not only an abundance of the very best button material 
but also a plentitude of pearls, and thus prove a source of much 
greater benefit than where mussels are reared for the shells alone. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 45 

DISCUSSION OF MUSSEL SPECIES. 

In the list of species here presented we have followed in most 
respects the classification and sequence given in Simpson's well- 
known Synopsis of the Naiades. In the spelling of the names, 
however, we have followed the suggestions of Lindahl a and have 
made also a few minor changes, such as the substitution of the older 
name Quadrula undata for Quadrula trigona, as suggested by Mr. 
Bryant Walker, and the transference of the Medionidus suhtentus 
(Say) to the genus Ptycliobranchus, and of Tritogonia tuberculata 
(Barnes) to Quadrula under the name Quadrula tritogonia, as sug- 
gested by Dr. Ortmann. Many other changes have been proposed 
which will probably in the end prove justifiable. We have avoided 
making any shifts between Pleurobema and Quadrula, although sev- 
eral have been advocated which may be desirable. The fact that 
both Pleurobema sesopus, Quadrula obliqua, and another perplexing 
form which we have found may have glochidia in two, three, or all 
four gills indicates that these two genera should really be united. 

We are very favorably inclined toward the new classification pro- 
posed by Ortmann, 6 but its present state of incompleteness and the 
uncertain position of many species, as well as our own conclusions 
regarding Quadrula and Pleurobema, make it seem best at present 
to use the older and better known system with the few exceptions 
noted above. 

1. Truncilla triquetra R,afinesque. Snuffbox. 

This attractive little shell occurs only in the upper part of the river. In all we 
procured 21 specimens, 19 of which were obtained at Salt Lick Bar. In the autumn 
of 1910 Mr. Boepple found it at Indian Creek, Cloyds Landing, Albany Landing, 
and in the Obey River at Celina, Tenn. 

It is in all probability considerably more common and widely distributed in the 
river than our collections would indicate. But it does not seem to occur as far down 
the river as Half Pone Bar or at Clarksville; if it did a few examples would certainly 
have been taken among the great number of small mussels collected in that region 
in Jane. 

Truncilla triquetra is a small species, dwelling in the shallower water. On account 
of its small size it is rarely or never taken on the mussel dredge or rake, but must be 
gathered by hand. It has a handsome, strong and thick shell, but is too small to have 
any commercial value. All our examples are pretty well eroded at the umbones. 

2. Truncilla brevidens (Lea). 

This species was not found in the main river at all and only at three stations alto- 
gether. It was most abundant in the Big South Fork opposite Parkers Lake Station. 
Three examples were procured in the same fork 2 miles above Burnside and one in 
Beaver Creek. It is too small to have any commercial value. 

All the specimens found were dead, but some had been recently killed by muskrats, 
therefore nothing was learned concerning its habits. It appears to be a species 

a Lindahl, J.: Orthography of names of the Naiades, The Journal of the Cincinnati Society of Natural 
History, vol. xx, no. 5, art. vm. 

6 A monograph of the Najades of Pennsylvania, reprinted from the Memoirs of the Carnegie Museum, 
vol. 14, no. C, Feb. 15, 1911. 



46 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

occurring in moderate-sized, clear streams with a rocky bottom, avoiding the smaller 
tributaries. 

3. Truncilla arcseformis (Lea). 

Rare; only one example obtained. This was procured in the Big South Fork 2 
miles above Burnside, Ky., and is rather peculiar in shape. 

4. Truncilla sulcata (Lea). Pewee, cat's-claw. 

Although this species seems to be pretty well distributed along a considerable 
stretch of the river, we obtained only occasional examples here and there along shore. 
Mr. Boepple found one in Caney Fork. It can probably be procured in larger num- 
bers during low water. It is common enough to be pretty well known to the clammeiv, 
who call it "pewee" on account of its small size, or "cat's-claw" because of the 
peculiar clawlike structures on the marsupial expansion of the shell of the female. 

5. Truncilla haysiana Lea. 

Our collection of this species is rather small, but it is probably more common than 
the collection would indicate, as it is too small to bite on the crowfoot hook and is 
easily overlooked. Most of the examples collected had been killed and cleaned by 
muskrats. It is one of the handsomest of the Truncillas on account of its beautifully 
polished epidermis, and it has an unusually thick and solid shell for the genus. It is, 
however, too small for manufacturing purposes. 

6. Truncilla capsseformis (Lea). 

Fairly abundant in the Big South Fork, where nearly all. the specimens had been 
killed by muskrats; in the main river we found it sparingly. Our shells are pretty 
badly eroded, very thin and brittle, with the marsupial expansion colored a dark 
green. The species is of no value for manufacturing purposes, being too small and 
thin. 

7. Truncilla florentina (Lea). 

Rare; the only specimen obtained was the dead shell of an old and very inflated 
female at Half Pone bar. In the autumn of 1910 Mr. Boepple found a specimen at 
Indian Creek bar. During low water probably many more could be obtained. 

8. Truncilla walkeri, new species. (See fig. 1, frontispiece.) 

A fine, large Truncilla with a honey-yellow epidermis and numerous capillar^ rays. 
Shell rather thin, elliptical in outline, much inflated in the females, only moderately 
in the. males. Anterior margin projecting and evenly rounded, ventral margin 
strongly convex in the larger males, much less so in the females and smaller males; 
posterior margin oblique, but usually well rounded in both sexes; dorsal margin 
comparatively long, straight, or slightly curved. Umbones narrow and flattened. 
Anterior, lateral, and posterior slopes all well rounded; umboidal ridge flattened and 
indistinct, especially in the females. In front of this ridge the males have a broad 
and shallow sulcus; in the females the marsupial expansion is very pronounced, and 
is usually limited anteriorly and posteriorly by a deep and narrow sulcus. It is 
somewhat like that of capsseformis, but is considerably swollen, especially in the 
larger females, instead of being flattened, and does not project as strongly. Lines of 
growth smooth, distinct, and close together. Ligament long, thin, and light brown. 

Interior: Pseudo cardinals large and thick, rather blunt and only slightly serrate or 
smooth; laterals long, high, thick, and slightly curved; anterior adductor scar slightly 
longer than wide, squarely truncated posteriorly ; posterior scar large, deeply impressed , 
and squarely truncated anteriorly much as in brevidens; pallial impression fairly dis- 
tinct, nacre milky white, thinner and quite iridescent posteriorly. 

This species was quite abundant just below the ford of the East Fork of Stones 
River near Walterville, Term. We found here 140 shells, most of them on shore and 
recently killed by muskrats, and 1 or 2 living mussels. The smallest specimen (male) 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 47 

measures 23.2 mm. long, 15.3 mm. high, and 8.8 mm. in diameter, the smallest female 
31 mm. long, 19.9 mm. high, and 12.9 mm. in diameter. The largest male measures 

57.7 mm. long, 42.7 mm. high, and 26.4 mm. in diameter, and the largest female 

52.8 mm. long, 39 mm. high, and 23.9 mm. in diameter. There are 49 females, the 
others being males. 

Walkeri, to Mr. Bryant Walker, one of our most eminent conchologists. 

9. Lampsilis ventricosa (Barnes). Pocketbook. 

Typical specimens of this species were obtained in two tributaries of the Cumber- 
land, Harpeth River near its mouth, and Stones River, in the east fork at Walterhill 
and the west fork at Murfreesboro, Tenn. Just how common or widely distributed it 
is in the streams above mentioned is not known. The examples found were excep- 
tionally fine and would make very good button shells. 

In the main Cumberland L. ventricosa seems to be quite rare, its place being usually 
taken by the closely related L. ovata. Indeed, the distribution and relationships of 
ovata and ventricosa as found in the Cumberland and its tributaries are exceedingly 
perplexing. A few examples found near Clarksville, and a dwarf shell found at Elk 
Creek shoals above Dover, however, offered exceptional difficulties in classification, 
fitting in neither with ventricosa nor ovata; the male shell would perhaps fall on the 
ovata side of the dividing line; the females on the ventricosa side, if indeed not rather 
beyond the limits of the typical shell; these shells, both male and female, were rather 
too thick and solid for ovata. A marked feature of those at hand is a deep pink tinge 
of the nacre posteriorly, "this tinge being pretty sharply limited to the posterio-dorsal 
area, a feature not common with either ventricosa or ovata. 

The female shells are considerably more inflated than the males and have a peculiar 
flattening of the lower part of the posterior margin. One of the female shells wa3 
sent to Mr. Bryant Walker, who remarks concerning it as follows: "No. 5456 is a 
most remarkable shell. I have never seen a female ovata with such an enormous 
expansion. Ventricosa not uncommonly tends that way, but not to such an extent. 
* * * This shell is comparable only with satur. [A variety of ventricosa, accord- 
ing to Simpson; satur is L. excavata, according to Frierson.] This shell is either an 
extraordinary abnormality of ovata or is ventricosa. In view of the occurrence of 
ventricosa both in the Harpeth and Stones, I am inclined to refer it to the latter." 

10. Lampsilis ovata (Say). Southern pocketbook; "grandma." 

A fairly common species throughout the entire length of the Cumberland, more 
numerous in the upper portions and upper tributaries. 

This species is one of the very few found in the Cumberland above the falls. Mr. 
Boepple obtained it at Pineville and Williamsburg and we found a few in the vicinity 
of the latter place and several examples just above the falls. Just below the falls it 
was abundant and common at the stations farther down. Associated with the typical 
form, which is relatively uncommon, is an aberrant form, more closely resembling 
ventricosa. 

The specimens of this aberrant form were at first identified as L. subovatus Say, 
described and figured by Call. a On examination of the literature, however, there is 
no " Unio subovatus Say/' and the name in Call's report is plainly a misprint for 
ovatus, the Unio subovatus Lea being an entirely different thing. 

Say's original description of "Unio ovatus" is brief and the figure poor, but recog- 
nizable; it is probably better known from Conrad's description and excellent figure. & 

The greater number of our specimens, however, differ considerably from the typical 
form. Beginning with the shapely, high-ridged clear yellow shell, which represents 

a Mollusca of Indiana, Twenty-fourth Annual Report of Geology and Natural Resources of Indiana, 
p. 481, pi. 39. 
* Conrad, Monography, p. 4, pi. 2. 



48 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

the species in its perfection, we have stained horn-colored examples, then deep- 
brown specimens and specimens with broad distinct rays. Inflated females are likely 
to have the ridge characteristic of ovata less markedly developed than males; in both 
sexes, however, there is a tendency for it to appear in all degrees of imperfect develop- 
ment until in some examples it is barely discernible. Indeed in one of our examples 
it is almost wholly absent, and we have a shell that, with the exception of purely 
individual features, can not be distinguished from a specimen of L. ventricosa from the 
upper Mississippi River. 

Judging from the soft parts of a single gravid female examined, the bodies of ventri- 
cosa and ovata are quite unlike, the mantle flap of ovata showing a peculiar mottling 
quite different from the markings observed in the other species. 

The variously modified forms of ovata are not only more abundant, but also more 
widely distributed than the type form. Just below Cumberland Falls most of the 
shells of this species are of medium size or smaller, nearly all are smoky brown, and 
several are well-rayed. At Indian Creek Bar brown and few-rayed individuals occur 
along with the typical form. At Goodall Island we found one with numerous distinct 
rays. 

The shells of the Rock Castle River are different from the others and can be told 
almost at a glance. They are dark brown, longer and heavier than those of the other 
streams and have the posterior ridge rather low; ovata takes nearly the same place in 
the Cumberland that ventricosa does in the upper Mississippi. 

We have always found ovata considerably inferior to ventricosa as a button shell, 
being thinner, smaller, and more brittle. The Rock Castle River ovata could possibly 
be used for buttons, but would furnish rather poor material. In the Cumberland the 
ovata is a rather valueless shell. 

Call's experience with ovata is different. He says it is "one of the largest that are 
found in American waters; * * * it also attains a much greater size than Barnes's 
form (ventricosa)." This may be perfectly true for some rivers, as shells vary greatly 
in size and thickness in different streams. 

11. Lampsilis multiradiata (Lea). 

Rather rare in the main river and found almost entirely in its upper portion. 
Occurs typically in small, clear streams and often in lakes. It is more common in 
the tributaries than in the main river. The specimens from both forks of Stones River 
are beautiful shells, typical in form, not much eroded, and with a clear, white nacre. 
The specimens from Rock Castle River, Big South Fork, and the main stream depart 
more or less from the typical form, being unusually elongate and sharp-pointed posteri- 
orly, rather thin, considerably eroded, and more or less stained or diseased in the 
n&cre or in the teeth. On comparing the Red and Stones River shells with the others, 
a marked difference was noted in the cardinal teeth. In these typical specimens 
the large posterior cusp of the right valve pointed more or less anteriorly, while in the 
Rock Castle River specimens and most of the others it pointed more or less poste- 
riorly. A few shells with intermediate characters in this respect were found, however. 

12. Lampsilis ligamentina (Lamarck). Mucket. 

This species is represented in the Cumberland chiefly by the southern mucket, 
Lampsilis ligamentina gibba Simpson, which differs from the typical form in being 
shorter and more compressed. The two forms grade into each other so impercep- 
tibly that it is impossible to find the point of separation between them. In the 
lower part of the river it approaches more nearly the typical form. 

The shell of the subspecies often has the epidermis more highly polished than in 
the type form, the nacre has more luster, and the valves are flatter and more uniform 
in thickness. The shells are therefore superior to those of the common mucket for 
manufacturing purposes and are sought after by shell buyers. Like the other forms 
of this genus this mucket carries its young in the gills through the winter. The 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 49 

glochidia fasten readily to our common spiny-rayed fishes. Some gravid examples 
of this form were collected by Mr. Boepple in the autumn of 1910 from the upper 
Cumberland and sent to the biological station at Fairport, and though the mussels 
were dead the glochidia were still alive and attached themselves readily to fishes. 

This is the most desirable form with which to stock the river and extensive plant- 
ings from the falls to the mouth would greatly increase its value as a mussel stream. 

13. Lampsilis orbiculata (Hildreth). 

Fairly common in the middle portion of the river, usually from 1 to 3 examples 
being found on each bed. 

We were struck with the remarkable similarity between this species and the south- 
ern mucket, Lampsilis ligamentina gibba. About the only way to distinguish between 
them was by the bright orange shade of the epidermis, and usually orange tint of 
the nacre of orbiculata, and it is easy to understand Call's a remark that Dr. Hil- 
dreth and the earlier naturalists seem to have considered this shell as a variety of 
Unio crassus Say (= Unio ligamentinus Lamarck, short and thick variety found in the 
Ohio), but Call adds: "It certainly would seem to be a good species." Our own 
studies and comparisons showed them more distinct than appeared at first glance. 
The difference is most plainly seen in the female shells, which differ considerably 
from the males, being truncate posteriorly and short and well swollen postbasally. 
They are well represented by Say's & figure of Unio abruptus, and look somewhat 
like a compromise between the southern mucket and L. ventricosa. Ortmann e 
says that this species "is not at all related to L. ligamentina as Simpson thinks; but 
it belongs to the ventricosa group of Lampsilis, for it has a well-developed flap on 
the mantle edge." 

L. orbiculata also very closely resembles L. higginsii which is more generally north- 
ern in its distribution, but the males of higginsii are shorter, more closely approach- 
ing Obovaria ellipsis. Orbiculata and higginsii are probably closely related. 

This is a very good button species, but so uncommon that it is not much of an item 
in the trade. 

14. Lampsilis Ueniata (Conrad). 

Rare; none at all were found in the Cumberland or in any of the tributaries except 
Stones River. It appears to be a species of small clear streams, and was found in 
the fine gravel at the edge of the water among the water- willows. 

15. Lampsilis picta (Lea). Painted mussel. 

Rather rare, and not taken by us in the main river. We found three in the Rock 
Castle River a few miles back from the Cumberland. Mr. Boepple, in the autumn 
of 1910, obtained it in the Big South Fork at Sloans Shoals, near Burnside. It is 
too small and thin to have any commercial value. Our largest example measures 
61 mm. long, 33 mm. high, and 17.5 mm. in diameter. 

16. Lampsilis punctoM (Lea). Spotted mussel. 

It is very like L. picta in color and outline, but differs in being more inflated and 
in carrying its thickness to the edge, so that its ventral margin is rather rounded and 
blunt, while that of picta is sharp. Both species are new to our collection. Mr. 
Bryant Walker, who identified them for us, called attention to the differences. The 
shell is thick anteriorly, but thins out rapidly behind the center. It has no commer- 
cial value on account of its small size, and most of our specimens are also badly eroded. 

17. Lampsilis perdix (Lea). 

Abundant in the Cumberland just below the falls. Mr. Boepple in 1910 found it 
as far down as Rowena and in the Obey River at Celina. It is common in Rock 
Castle and frequent in the Big South Fork. 

a Mollusca of Indiana, Indiana Geological Report, p. 493. 
l> American Conchology, pi. 17. 
c Nautilus, vol. xxm, no. 9, p. 119. 



50 MUSSELS OF CUMBERLAND EIVEE AND TRIBUTARIES. 

This species bears a general resemblance to an elongate flattish L. ligamentina. 
Unstained shells are easily recognized by the character of the rays, which are broken 
up and more strongly marked in places, making a series of heavy green blotches. 
Another peculiarity is the short lateral teeth, 1 in the right valve and 2 in the left; 
these are low and blunt, and separated from the cardinals by a wide interspace. 
Our older shells are badly eroded and so stained and discolored that the character- 
istic rays and blotches are absent. Such specimens can be recognized by the narrow 
border of latest formed nacre, which is yellowish or reddish and semitranslucent. 
Our shells usually have the nacre badly stained. Even if obtained free from stains 
they would make rather poor button shells, as they are somewhat brittle. In thick- 
ness they are about equal to a thin mucket. A few of the examples have brick-red 
pimply patches on the interior which probably indicate the presence of parasitic 
trematodes. No parasites, however, were noted. 

18. Lampsilis anodontoides (Lea). Yellow sand-shell. 

Rather uncommon, distributed chiefly through the central portion of the river, 
and never forming a large percentage of any of the beds. This species thrives best 
on sand bars in rather shallow water. It is generally confined to large streams. It 
is one of the most active of the mussels, responding quickly to changes in environment 
by moving about. This is by far the most valuable of the fresh- water mussels, the 
shells being generally used for export and in the manufacture of knife handles. 

This species is easily propagated, the glochidia fastening readily to most of the 
common spiny-rayed fishes, such as sunfishes, bass, etc. On June 13 we found a 
number of gravid shells at Meeks Bar. Some sunfishes were caught, a tub was pro- 
cured, and an infection made. The infected fishes were then liberated into the Cum- 
berland in front of the blank factory at Clarksville. 

19. Lampsilis fallaciosa (Smith). Slough sand-shell. 

Rare in the Cumberland and not found in any of the tributaries. This species 
thrives best along shore in shallow water with a rather lively current and muddy 
bottom. Such conditions exist only in the very lowest portion of the Cumberland. 
From Kuttawa to the Ohio side sloughs are more common and the species is probably 
more abundant. The nacre of most specimens secured is stained. This is a first- 
class species for the manufacture of buttons, but it would be unprofitable to plant in 
the Cumberland because of the absence of favorable locations for its best development. 

20. Lampsilis recta (Lamarck). Black sand-shell. 

Rather common throughout the entire length of the river, but nowhere abundant. 
Many of the shells are badly eroded and stained; none are deep pink throughout, 
but are pale pink about the cardinal teeth and in the umbonal cavity. 

Good white-nacred shells of this species are exceptionally excellent button shells, 
and where select stock could be obtained would be one of the most desirable species 
to propagate. 

21. Lampsilis lienosa (Conrad). 

The specimens we have are hardly typical and were with some doubt identified as 
this species. It is a small species of no commercial importance. 

22. Lampsilis vanuxamensis (Lea). 

The females of this species were gravid June 6. They are peculiar in having the 
marsupial expansion of the shell rather limited in area, not extending to the posterior 
end, but followed by a pointed extremity. In this localization of the shell they 
remind one somewhat of the Truncillas. The shells are small, red nacred, and of 
no value. 

23. Lampsilis trabalis (Conrad). 

Found only in the upper part of the river and its tributaries. The females are not 
markedly swollen posteriorly, but differ from the males in being shorter and broader. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 51 

Nearly all we found were dead shells, usually badly eroded at the umbones. Mr. 
Boepple found the species as far down as Cloyds Landing and in the Obey River at 
Celina. As found, the epidermis is generally jet black, usually due to the shells 
being stained. On being cleaned with acids they exhibit beautiful rays. This is a 
small species of no commercial importance. 

24. Lampsilis parva (Barnes). 

Rare; none at all in the Cumberland; indeed it has not been reported from that 
river. But we obtained one specimen in the East Fork of Stones River at Water- 
ville. This was a slender shell; length 27 mm., height 15 mm., width 11 mm. Nacre 
beautifully white and iridescent. 

25. Lampsilis glans (Lea). 

Rare; none at all in the Cumberland; 10 specimens from the tributaries. Those 
found were in gravel in shallow and rather swift water. In general it prefers quiet 
streams with muddy banks and burrows in the firm mud. It is also frequently found 
in lakes. 

One of the smallest of our species; too small for commercial use, and with a rich 
purple nacre. In one specimen, a female found in Roaring River, the peculiar 
glands of the mantle, small white cylindrical objects on each side, were protruded 
and were undergoing spasmodic movements. 

26. Lampsilis alata (Say). Pancake; pink hatchet-back. 

While not a rare species in the Cumberland, this is not especially common. In a 
few of the beds it is entirely absent, and in many only one or two shells were found. 
It never exceeded 4 per cent of the catch of any of the beds, and is usually less than one. 
It is well distributed throughout the entire river. It prefers rather deep water and 
a soft, muddy bottom. The shell, on account of its thinness and red nacre, is of no 
value whatever. 

27. Lampsilis gracilis (Barnes). Paper-shell. 

Frequent enough to be a rather familiar species among clammers, but not so abun- 
dant as to be a nuisance. It has much the same distribution as alata, but is less common. 
We usually obtained only 1 or 2 from a bed. Our shells are rather badly worn at the 
umbones. As this thin-shelled species is of no value whatever, but readily catches 
the mussel hooks, it proves to be a nuisance when present in large numbers where 
clamming operations are being carried on. 

28. Lampsilis Isevissima (Lea). Paper-shell. 

Rare; only one specimen found in the Cumberland; this was at Meeks Spring Bar. 
It seemed to be more common in the Harpeth. This species closely resembles L. 
gracilis in general appearance, but has, among other distinguishing features, a beauti- 
fully polished epidermis. Our examples have a number of peculiar rays, consisting 
not of a different pigmentation of the epidermis but of a series of short, finely wrinkled 
lines. 

29. Lampsilis leptodon Rafinesque. 

This fragile, thin-shelled species is rare in the Cumberland. The only examples 
obtained were collected by Mr. Boepple at Albany and Cloyds Landing in the autumn 
of 1910. 

30. Medionidus conradicus (Lea). 

This species is confined chiefly to small streams. It is exceedingly abundant in 
the Rock Castle River at Livingston, Ky., the sandy bottom being almost covered 
with these animals, which showed up as narrow black lines, the mantle and exhalent 
and inhalent apertures being thin and black. It is also abundant in Roaring River. 

In the Cumberland we found it just below the falls and at Salt Lick Island. All 
the shells were badly stained and eroded, and for this reason, as well as on account 
of its small size, it has no commercial value. 



52 MUSSELS OP CUMBERLAND RIVER AND TRIBUTARIES. 

31. Obovaria retusa (Lamarck). Golf-stick. 

Although we obtained only a few specimens of this species, scattered valves were 
frequently found along shore, and there is reason to believe that it is considerably 
more common than our small collection would indicate, although by no means 
abundant anywhere. In the Cumberland it attains a rather large size, our largest 
shell measuring 68.5 mm. long, 74.5 mm. high, and 46.9 mm. in diameter. It is a 
heavy and solid shell, but the deep purple of that portion of the nacre within the 
pallial line makes it valueless for buttons. All our shells are somewhat eroded at 
the umbones. Two of them are considerably less retuse than the others, somewhat 
approaching 0. circulus in this respect. All have the epidermis somewhat paler 
posteriorly, but not so markedly so as is usually the case with 0. circulus. 

32. Obovaria circulus (Lea). 

Rather common in the main river from Burnside to Half Pone Bar. This species 
produces too small a shell to be of much importance to the button trade. The larger 
shells would furnish two or four blanks apiece, and are excellent both as to material 
and thickness. The nacre seems to be unusually durable and retains its firmness and 
luster long after others have become chalky. 

33. Obovaria ellipsis (Lea). Missouri niggerhead. 

This species is chiefly northern in its distribution and does not attain large size in 
the Cumberland. Although in its shell characters it bears considerable resemblance 
to some of the Quadrulas, especially the niggerhead, Q. ebena, it is really more closely 
related to the sand-shells. Where it attains large size it is an excellent button shell 
and would be a fine species to propagate, but the reduced size of the shell in the 
Cumberland indicates that the conditions there are not favorable. We found gravid 
examples above Clarksville early in June. 

34. Plagiola securis (Lea). Butterfly. 

This species is fairly common throughout the entire length of the river below the 
falls, and, while not abundant enough to make a large percentage of the shells taken for 
commercial purposes, it makes a fair sprinkling in most of the clammers' piles. It 
seems to thrive exceptionally well in the Cumberland and is more common here than 
in most rivers. The shell, especially of young to medium-sized, well preserved males, 
is one of the most attractive among the Unionidse. In the Cumberland there is a 
marked difference between the shells of the males and females, that of the former being 
flat and compressed and of rather uniform thickness, while those of the females are 
much more tumid and swollen . The measurements of a fairly typical male (F5086) 
of medium size are 54 mm. long, 44 mm. high, and 21.1 mm. in diameter, while those 
of a tumid female of about the same length (F2660) are 55.3 mm. long, 45 mm. high, 
and 33.7 mm. in diameter. In the lower part of the river the nacre is somewhat 
spotted, but upstream the shells are free from stain. On account of its excellent 
luster, flatness, and uniform thickness, this is an excellent button shell, the males 
being much superior to the females. 

Females were found gravid May 29, and were in the height of the breeding season 
from about June 3 to 16. This would be a very valuable species with which to stock 
the river. 

35. Plagiola elegans (Lea). Deer-toe. 

This species is not as common nor as widely distributed as the preceding. Large 
shells can be used in the manufacture of buttons, but the great majority are too small. 
The largest example found was a single valve 59 mm. long, picked up at the foot of 
Gowera Island. The beautifully tesselated green markings on the epidermis make 
it an attractive shell when perfect. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 53 

36. Plagiola donaciformis (Lea). 

This dainty little species is more limited in its distribution in the Cumberland 
than either of its two relatives. A peculiarity of the species at Half Pone Bar was the 
frequent unfolding of the anterior ventral portion of the shell, the inner layer being 
folded back against the rest, as if by some injury. The specimens found here were 
unusually thin-shelled and frequently had the nacre well tinged with pink. Perfect 
specimens of this shell are among the most attractive to be found in the Unionidse, but 
the Cumberland examples, especially those from Half Pone Bar, are badly worn at 
the umbones, so that even small specimens have the appearance of age. This is one of 
the smallest of the mussels — too small to be of any use for manufacturing purposes. 

37. Cy progenia irrorata (Lea) . 

This species is of rather infrequent occurrence in the Cumberland. We found none 
at all in any of the tributaries, and usually found only one or two on each bed exam- 
ined. The species seems to inhabit rather deep water, since we never saw any crawl- 
ing around on the shallow bars. Most of the examples are rather small, and some have 
a shallow sulcus running over the middle of the disk from the umbonal region to the 
postventral margin. 

A very solid shell, but of little commercial value, as it is rather brittle and has pink 
tips. The few shells that get into the clammers' piles are generally worked up, 
however. 

38. Obliquaria reflexa (Rafinesque). Three-horned warty-back. 

One of the most common shells of the river, and found throughout its entire extent. 
Although a rather small shell, this is so thick and solid that it is used to a considerable 
extent in the manufacture of buttons, each valve furnishing one or two small blanks. 
The species has a long breeding season, spawning through almost the entire summer, 
the young being extruded in white cylindrical masses. Some of these spawn masses 
were seen lying on the gravel at Half Pone Bar June 16. Shells of females are some- 
what fuller anteriorly than the males and can usually be distinguished after some 
practice. The Cumberland specimens are not so beautifully rayed as those from the 
upper Mississippi. 

39. Ptychobranchus phaseolus Hildreth. Kidney-shell. 

Scattered in the upper Cumberland from the falls down to Half Pone Bar. Although 
this is a species of rather wide distribution, especially southward, and is by no means 
a rare shell, it is never found in great numbers or making a large per cent in any bed. 
The clammer rarely gets over a half dozen or dozen to the ton; the nacre is white, 
with a soft satiny luster; the shape is nearly that of Unio gibbosus, and the species 
would probably make a fair button shell. 

40. Ptychobranchus subttntus (Say). Fluted kidney-shell. 

This species in Simpson's Synopsis is placed in the genus Medionidus. Dr. Ort- 
mann, however, has removed it to Ptychobranchus, and, although we have seen no 
gravid examples, we are inclined to follow him in this regard on account of the close 
resemblance of the shell to that of P. phaseolus, differing from that species chiefly in 
its thinner shell, greater inflation, and the presence of costse on its posterior slope. 
On account of its small size and its thinness it has no commercial value. 

41. Dromus dromas (Lea). Dromedary mussel. 

In the main river this shell is of occasional occurrence from Mill Springs Bar, in the 
upper river, down to Red Rock Bar, below Clarksville, Tenn. We usually obtained 
one or two specimens at a station. The shells are rather heavy and inflated, though 
the hump on the disk, which is characteristic of the species, is not nearly as prominent 
as in some specimens from the Washington collection obtained by Mr. Boepple in the 
Clinch and Holston Rivers. Some of the shells are beautifully rayed, especially 



54 MUSSELS OP CUMBERLAND RIVER AND TRIBUTARIES. 

anteriorly, but the greater number are too deeply stained for the rays to show. In 
the living animal the mantle is prettily rayed. 

The shape, size, and solidity of the shell of this species make it suitable for the 
manufacture of buttons, but unfortunately it is too brittle and hard, resembling 
Pleurobema sesopus in this respect. About one-third of the shell, moreover (the tip 
part), is of a pink tinge, which runs entirely through the shell, making it of no value. 

42. Dromus caperatus (Lea). Fan mussel. 

The examples of Dromus obtained in the Big South Fork of the Cumberland differ 
from those found in the main river by being considerably flatter, with the hump on 
the disk less pronounced or nearly absent. These flattened shells represent the species 
caperatus (Lea). Our series indicate that the two forms run together. In young 
specimens, before the step-off is formed, it is doubtful if dromas and caperatus could 
be distinguished. 

From what has been said concerning the relationship between this and the pre- 
ceding species it may be readily inferred that this species also, from a commercial 
standpoint, is valueless. 

43. Strophitus edentulus (Lea). Squaw-foot. 

We found only a few examples of this species. It has a fragile shell, which disin- 
tegrates quickly and is probably more common than our small collection would indi- 
cate. Mr. Boepple found it at, Pineville, the highest point at which the river was 
examined. It is a species which occurs in all sorts of situations — in both small and 
large streams and in lakes. Two of our specimens have a pink-purple nacre; in the 
others it is of a yellowish cast. The species is of no value on account of its thin, brittle 
shell. It is exceedingly variable, and presents many puzzling forms. According to 
Mr. Bryant Walker our specimens represent the form shaefferiana Lea. 

44. Anodonta imbecillis (Say). 

The distribution of this fragile, beautiful species is almost identical with that of 
A. grandis. Of the two found in Haynes Lake one was gravid (Sept. 3). The glo- 
chidia are rather large, chestnut-shaped in outline, brown, and fill the entire outer 
gills. The species remains gravid through the winter. The Haynes Lake shells 
contained several Atax apiece. 

45. Anodonta grandis (Say) . 

This species was not found in the main river. In general, conditions throughout 
the whole Cumberland system are not favorable to its development. The small 
tributaries are too swift and rocky, and the Cumberland itself is lacking in the quiet, 
muddy sloughs in which A. grandis can thrive. The only river examples we found 
were in the Stones River, a few in the East Fork near Walterhill, Tenn., and several 
in the West Fork near Murfreesboro. At the last-mentioned place it had apparently 
once been abundant in the vicinity of the railroad bridge, where it had thriven in the 
mud of the deep, quiet pools among the water- willows. A number of shells, recently 
killed by pearlers, were lying on the bank. These were large, heavy shells, unusually 
thick for the species, and varied considerably in shape, some of them being markedly 
elongate. 

In Haynes Lake, a shallow, muddy pond below Clarksville, Anodonta grandis was 
fairly abundant, and about 30 examples were secured. These were more shapely, 
of a larger size than those from Stones River, and much thinner. They are indeed 
the largest and finest examples of the species we have ever seen and represent the 
form gigantea Lea. The largest example measured 201.3 mm. long, 112.5 mm. high., 
and 82.3 mm. in diameter. These shells are peculiar in having two distinct colors of 
nacre, about half of them being dark purple, while the other half are a beautiful, 
lustrous, creamy white. The reason for this difference is not apparent; parasites are 
almost entirely absent. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 55 

46. Lastena lata (Rafinesque). 

Very few examples seen in addition to those enumerated in the table. Mr. Boepple 
obtained it at Burnside, Albany Landing, and Cloyds. Its apparent scarcity is due 
in part to its habits. It can not be caught on the crowfoot hook, but must be obtained 
by wading, and is best secured when the water is low and clear. The species appears 
to prefer gravel bars with a rather swift current. The shell is beautifully polished 
and rayed, and is very thin, cracking easily when exposed to the air. Our examples 
are rather badly eroded. 

47. Anodontoides ferussacianus (Lea). 

Rare; only a few specimens found. A thin, fragile Anodonta-like shell of no com- 
mercial value. 

48. Pegias fabula (Lea). 

A rare species of which we found only two living and four dead specimens in the 
Rock Castle River near Livingston, Ky. They are quite small, the smallest measur- 
ing 22.7 mm. long, 15.5 mm. high, and 11 mm. in diameter, and the largest 31 mm. 
long, 20 mm. high, and 14 mm. in diameter. In their perfect condition these must 
be very attractive little shells, but our specimens are very badly eroded. 

49. Symphynota costata (Rafinesque). Fluted shell. 

Occasional in the upper Cumberland from the falls down to the foot of Gowers 
Island. Occurs typically in moderately small streams and appears to be entirely 
absent from the lower stretches of the Cumberland. It is rather common in the 
various tributaries. The Stones River shells were exceptionally thick and heavy, 
and bore a goodly number of dead or soft pearls. 

On account of its yellow nacre and tendency to crack this species is of no use in the 
manufacture of buttons. 

Several of our specimens have numerous deep wrinkles extending ventrally over 
the posterior half of the disk. One is unusually shortened, truncate posteriorly and 
produced forward, and has well-marked rays, while another medium-sized shell from 
a mile below the falls is unusually elongate. 

50. Symphynota complanata (Barnes) . White heel-splitter. 

Rare; only two examples of this species were found in the entire Cumberland. 
The shells were small, thin, and badly stained. These were obtained on Red Rock 
bar below Clarksville. Fragments of large strong shells were found in the Harpeth 
River. This species thrives in a muddy bottom and is often found in sloughs. Under 
especially favorable conditions it produces a fairly thick large shell which furnishes 
usable button material, but the Cumberland shells of this species have no value. 

51. Alasmidonta minor Lea. 

Confined to the upper river and tributaries. So far as our experience goes, this 
species is found typically in small streams, living in the sand between rocks. It may 
live along the border of large streams, but on account of its small size would be easily 
overlooked. Most of the specimens found had been killed by muskrats. The shells 
were all badly eroded and so deeply stained that the characteristic rays were ob- 
scured and the nacre rather badly stained. 

This species is always too small to have any commercial value. Our smallest 
example measures 17 mm. long, 11 mm. wide, and 6 mm. in diameter, and our largest 
45 mm. long, 28 mm. wide, and 18 mm. in diameter. 

This species closely resembles A. calceola, a better known and more widely dis- 
tributed species, but has a heavier shell and teeth and darker epidermis, and is some- 
what flatter and longer. 



56 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

52. Alasmidonta truncata B. H. Wright. Elk-toe. 

This is not a common mussel in the Cumberland and is, generally speaking, a spe- 
cies of rather small streams and the upper courses of larger rivers. All the shells 
found were dwarfed, very thin and eroded, and with the epidermis rather badly 
stained. 

When well developed this is an attractive shell, but it is always too thin and fragile 
to have any commercial value. 

According to Mr. Bryant Walker, there is no difference between this and A. margi- 
nata Say, and our thin dwarf specimens lend probability to this view. As we have 
seen but few marginata we have no means of comparing them. As Simpson has sepa- 
rated the two forms, however, and ours are within the geographic range of truncata, 
we retain for the present Simpson's name. 

53. Margaritana mpnodonta (Say) . Spectacle case. 

Occasional from Snows Island, where we first encountered it, as far down as Dover 
and perhaps beyond. The shells are fragile and break and crack easily, and disap- 
pear soon «iter dying. The species has no commercial value. 

54. Unio gibbosus Barnes. Lady-finger; spike. 

Unlike Unio crassidens this species is not especially abundant in the Cumberland. 
Though distributed throughout the entire length of the river, at many stations only 
a half dozen specimens were found, and nowhere did it rise above 4 per cent of the 
entire catch. In the Cumberland above the falls it is about the only species found. 
In the Clear Fork at Jellico, Tenn., and Savoy, Ky., it was abundant, forming about 
90 per cent or more of the entire mussel population, and numerous dead shells recently 
killed by muskrats were found along shore and at the base of the water-willows. 

These Clear Fork examples were all small dwarf shells with a rather pale nacre. 
They approach a well-marked form found in Green River, Ky., and other southern 
streams. The Clear Fork flows through sandy and shaly country and the water may 
be too deficient in lime to promote good shell growth. Immediately below the falls 
we encountered the normal full-grown form which is the one of the main river. 

Gravid examples of this species were found during the entire summer. 

55. Unio crassidens Lamarck. Elephant-ear. 

Exceedingly abundant, especially in the upper part of the river. It is a species of 
large streams, and we did not find it in any of the tributaries nor above the falls. In 
the upper part of the river this shell is a decided nuisance, forming a large part of the 
clammer's catch, taking much of his time and labor and yielding little in return. It 
is generally known as the "pink," and clammers, on their prospecting cruises, note 
down the percentage of "pinks" and "whites," from which to judge the value of a 
bed. It is the great abundance of this species that makes the section of river from 
Burnside to Celina unprofitable clamming, and the problem of making this stretch a 
valuable clamming ground consists as much in the reduction of this species as in the 
increase of valuable kinds. 

U. crassidens exhibits considerable modification as one ascends the Cumberland. 
In the lower stretches of the river most of the shells are the rather elongate form, 
which seems to be most common the country over. As one advances upstream these 
elongate shells gradually give way to a short and chunky variety. 

The shells from Half Pone bar and a few from Mill Springs and Salt Lick bar show 
rather well-marked rays; most of the others are rayless. 

Occasionally shells with the na^re very pale or almost white are found. These are 
called "white-pinks" and are acceptable to the buyer. Even the more or less 
markedly pink ones are beginning to be used, but there is little demand for them and 
they always bring a rather low price. The shells work up exceptionally well, being 
soft and free from grit. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 57 

While at Clarksville we were informed that the superintendent of one of the smelting 
furnaces along the river had been trying cull shells as a flux and found them satisfactory. 
It is doubtful whether this utilization, however, will make an important market for 
them. 

56. Pleurobema clava (Lamarck). Club-shell. 

Generally rare, and not found at all below Burnside. The shells are all badly 
eroded and discolored; one of them is unusually elongate, and several show a rather 
well-marked, broad and shallow furrow in front of the posterior ridge. We have 
usually found this species most abundant in small streams, and this may explain its 
absence from the greater part of the Cumberland. It is a rather handsome shell but 
too small to have any commercial value. 

57. Pleurobema crudum (Lea). 

This species does not appear to be common or widely distributed. All our examples 
are rather small shells, somewhat resembling a much-flattened Quadrula subrotunda, 
but with the epidermis of a brighter yellow and the rays quite distinct, well defined, 
and broken up into blotches. 

58. Pleurobema sesopus (Green). Bullhead. 

We did not see many examples of this species in the Cumberland, but it is common 
enough to be well known among the clammers. In the upper Mississippi it is called 
"bullhead" or "sheepnose," and is used in button manufacture, although it is ranked 
as a rather low-grade shell on account of its brittleness. In the Cumberland it is so 
hard and flinty that no attempt at all is made to cut it as it breaks saws. The clammers 
call it "clear profit" because they are "the only ones who get anything out of it." 
A small example obtained at Half Pone bar was of a beautiful yellow color; the older 
ones are brown. 

The systematic position of this species is in doubt. It seems to stand between 
Quadrula and Pleurobema. Simpson <* was not certain as to where to place it, having 
seen only one example gravid, and it with the gills partly filled. At the biological 
station at Fairport one was found with only the inner gills filled with glochidia and 
another with all four. Sterki & has found glochidia in all four gills. Usually, how- 
ever, only the outer gills are used as a marsupium. 

59. Quadrula tritogonia (Barnes). Buckhorn; pistol grip. 

This is the Tritogonia tuberculata of Simpson's Synopsis. At the time the Synopsis 
was written the gravid female was not known. The shell stood pretty much by itself, 
and Mr. Simpson, who was struck by certain peculiar features, especially the note- 
worthy difference between the male and female shells, formed a separate genus for it. 
Since the discovery by various students that it bears young in all four gills, there is a 
general tendency to place it in the genus Quadrula, and Dr. Ortmann, who was the first 
to propose the shift, suggested the name given above. The species is quite aberrant; 
none of the other Quadrulas resemble it very closely, the nearest approach being some 
of the elongate Quadrulas such as q/lindrica, especially the rough subspecies strigillata 
or Quadrula trapezoides from the south. The marked difference between the males 
and females is unique among any related forms and entitles it at least to subgeneric 
rank. 

This species is not rare in the Cumberland and was obtained in small numbers at 
most of the stations from the falls down to Dover. Our specimens are mostly of 
medium size and a number have the nacre rather badly stained. They exhibit 
but little variation among themselves or from the form as generally known. The 
nacre of all but two is white; in these two, obtained near Clarksville, it is pink. 

a Synopsis of the Naiades, Proceedings of United States National Museum, vol. xxn, p. 745 and 764. 
6 According to Ortmann, Nautilus, vol. xxn, no. 10, Feb., 1909, p. 100. 



58 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

Where it attains its best development, the buckhorn is an excellent button shell, 
indeed one of the best. It does not find the most favorable conditions for growth and 
development in the Cumberland, however. It is not as yet amenable to propaga- 
tion on a large scale, as it is but rarely that one finds gravid examples. 

60. Quadrula perplicata (Conrad). 

The plicate Quadrulas of the Cumberland, especially the middle portion of the 
river, are rather peculiar shells, lying somewhere between typical plicata and undu- 
lata. The beaks are too low and flattened for plicata and the shells are too heavy 
and a trifle too inflated for undulata. A marked feature .about them, in addition to 
their general rotundity of outline, is the fact that they usually taper to a point posteri- 
orly. The clammers call them the "round-lake," and say that in proper conditions 
they are good pearl bearers. The folds are few and gently rounded. Mr. Bryant 
Walker, who examined them, is of the opinion that they are perplicata. We obtained 
some good specimens at Meeks Spring bar. Our largest measures 119 mm. long, 86 
mm. high, and 56 mm. in diameter. At Half Pone bar a particularly interesting and 
instructive lot of young shells were obtained. These are inflated and rotund, 
approaching a spherical form with a greenish epidermis. Though quite small, they 
are so worn at the umbones that they look like old shells and no beak sculpture is 
shown. The smallest measures 17 mm. long, 15 mm. high, and 10 mm. in diameter. 
Farther up the river, at Cloyds Landing, this shell approaches undulata, while in 
Stones River, near Murfreesboro, the real undulata is found. 

The shells are thick, solid, and heavy, but the nacre is spotted and they form rather 
poor button material. If they could be obtained free from spots, they would have a 
good market value. 

61. Quadrula undulata (Barnes). Three-ridge or blue-point. 

Beautiful examples of this species are common in the West Fork of Stones River 
near Murfreesboro, Tenn. It is also found in the East Fork near Walterhill. The 
young examples are yellowish brown, well compressed, and entirely free from erosion, 
so that the umbones show the sculpture very plainly. This consists of four or five 
high, coarse ridges, the first-formed ones crescentic, the older ones gradually vanishing 
backward until the last one is a short, low tubercle. The undulations are deep and 
crossed by numerous small furrows. A noteworthy feature of these shells is the great 
distance of the pallial line from the margin. The shells are somewhat spotted, but 
the spots are small and they would yield a fair amount of good button material. 

62. Quadrula heros (Say). Washboard. 

This is a species of large rivers. It is not found in the upper part of the Cumberland, 
but is abundant in the lower river. The first we saw was at the Mill Springs bar. 

This species bears the largest and heaviest shell of the North American Unionida?. 
It becomes rather large in the Cumberland, but not as immense as in the Wabash and 
some parts of the upper Mississippi. Our largest shell measures 162.8 by 115 by 62.4 
mm. Our collection exhibits little variation. From the unusually large number of 
small examples seen it appears that the species is exceptionally prolific in the Cum- 
berland, especially about Half Pone bar and Owl Hollow bar above Clarksville. 
All our examples are somewhat eroded at the umbones, but only two or three badly. 
The young examples are noteworthy for having the finely waved broken sculptures, 
characteristic of the umbones of the older specimens, over the entire disk and the 
plications rudimentary or only faintly developed, so that they do not closely resemble 
the old. 

We found no gravid examples. They are indeed very rarely found, and nothing 
is known at present about its spawning habits or as to what fish acts as host to the 
embryos. a 

« Since the above was written investigators at the Biological Laboratory at Fairport have thrown, 
considerable light on the breeding habits, hosts, etc., of this species. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 59 

In some rivers, as parts of the Illinois, this shell does not become stained early, 
and the younger shells furnish excellent button material. For the common run of 
buttons this shell is becoming one of the most important species, as its large size and 
expanse allows it to be worked up readily into buttons of various sizes, and the stains 
can be bleached out or the buttons "smoked" or artificially dyed. In the Cumber- 
land the nacre becomes badly stained, even when the shell is quite small, and the 
washboards are always sorted out and sold separately as low-grade shells, bringing 
but $2 to $5 per ton when first-grade shells are bringing $6 to $8. 

But few parasites were found, and we have as yet no clue to the cause of the dis- 
colored spots on the nacre. These spots are usually circular in outline and frequently 
have what appears to be a foreign body in a small raised pustule at the center. The 
fresher stains, or those near the surface, do not really permeate the nacre, but are 
composed of a flat hornlike skin overlying it and can be softened by acids and scraped 
away from the unstained shell beneath. The older, duller stains are doubtless the 
same thing covered by layers of nacre. 

Many of our specimens are interesting as showing with unusual clearness the path, 
during growth, of the posterior adductor muscle scar, the anterior border of which is 
dimly defined, while straight converging lines from the dorsal and ventral borders 
of the scar lead up into the umbonal cavity. One of our specimens has a pinkish 
nacre. 

63. Quadrula cylindrica (Say) . Rabbit 's-foot. 

Occasional to abundant in the upper part of the river. On account of its narrow 
cylindrical shape it is of little value for buttons; the nacre, moreover, is frequently 
diseased and stained. The flesh is usually orange yellow and the gills, when filled 
with glochidia, markedly so. Some of our examples are well covered with small 
tubercules over the anterior portion of the disk, approaching the subspecies strigillata. 

This is a rather active species, the most active of the Quadrulas. Its elongate form, 
in which it differs markedly from its nearest relative, metanevra, and indeed from all 
Quadrulas in general, may be an adaptation to an active life. 

64. Quadrula metanevra (Rafinesque). Monkey-face. 

This well-known button species is fairly common. A few were to be found at nearly 
every station, clam pile, or mussel bed. It was not abundant enough, however, to 
form more than a sprinkling among the shell piles, and it cuts a rather small figure in 
the button industry of the Cumberland. On account of its luster and solidity it is 
very acceptable to the manufacturers. It would not be worth propagating, however, 
as there are plenty of better species. We found one example of this species gravid on 
the last of May. 

65. Quadrula tuberosa (Lea). 

Rare and collected only in the upper river. In the autum of 1910 Mr. Boepple 
obtained it at Sloans Shoals in the South Fork near Burnside, at Selfs Bar, and at 
Cloyds Landing. 

66. Quadrula fragosa Conrad. 

This species is occasional, and in some places abundant, in the lower Cumberland. 
It does not appear to "bite" readily on the crowfoot hook and the few examples 
taken by clammers are apparently no indication of its abundance. Small mussels 
of this species are a favorite food of the muskrat. Of a large pile of shells cleaned 
out by these rodents near Meeks Spring Bar, nearly all were this species and Obliquaria 
reflexa, although other mussels appeared to be common in the vicinity. 

This species is very similar to Quadrula lachrymosa (Lea) and the differences between 
the two are difficult to express either by description or figure. It is somewhat more 
square-cornered, more inflated, and the tubercles on the posterior slope are more 
markedly arranged in rows, forming costae. This species does not become as large as 



60 MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 

Q. lachrymosa and is of little commercial value. We found gravid examples below 
Kuttawa May 17 and at the foot of Dover Island May 29. All four gills serve as 
marsupia and are thick and pad -like. 

67. Quadrula pustulosa (Lea). Warty -back. 

Common throughout the entire length of the river. Our shells exhibit a marked 
uniformity in general appearance, being rather inflated with only a moderate num. 
ber of low tubercles. A few shells found a mile below Cumberland Falls are almost 
entirely smooth. With the exception of the Half Pone Bar specimens most of the 
shells have a cloth-like epidermis. 

The warty-backs of the Cumberland are as a rule rather undersized, and their 
inflated form is something of a disadvantage, so that they are not as valuable as in 
some other streams. 

68. Quadrula cooperiana (Lea). Cumberland pigtoe. 

Not rare in the Cumberland. The proportions of the shell vary considerably, some 
being higher than long and others longer than high. The older examples are gen- 
erally more elongate than the younger. The shells also vary somewhat as regards 
degree of inflation. One of the young shells has the epidermis faintly rayed, the 
others are eradiate. Three of the shells have the epidermis polished and shining; 
in the others it is dull. The nacre is sometimes a pale suffused pink within the 
pallial line, but in the majority of cases it is pure white. This is regarded as a very- 
fair button shell. In appearance it lies intermediate between pustulosa and granL 
fera. From granifera it can always be distinguished by the color of its nacre. It is 
usually longer and flatter than pustulosa, and there are peculiarities of epidermis, 
disposition of pustules, and shape of teetn that taken together help to separate them. 
They can always be separated if in the flesh, as cooperiana always has an orange- 
yellow flesh. The ova which fill the gills are bright yellow. 

We found only two examples gravid, early in June. The developing ova were 
borne in the outer gills and gave it a sulphur-yellow color. 

Dr. Ortmann removes this species from the genus Quadrula and places it in Pleu- 
robema; he remarks that it is closely related to P. sesopus. We are rather favorably 
inclined to this view, but in view of the fact that these two genera need a thorough 
revision and may possibly run into each other we prefer at present to leave it where 
Simpson placed it, among shells that it strongly resembles. 

69. Quadrula rubignosa (Lea) . Wabash pigtoe. 

This .species was found nowhere except in the East Fork of Stones River at Walter- 
hill, Tenn. The shells show very little difference in general appearance, except 
that in the smallest the posterior ridge is poorly defined, and one of the medium- 
sized examples is somewhat more rounded, and has a lower posterior ridge. Large 
examples of this species make a moderately good button shell. 

70. Quadrula undata (Barnes). Pigtoe. 

This, as Bryant Walker has shown, ° is the proper name for the Quadrula trigona 
(Lea) of Simpson's Synopsis. Ortmann t> regards it as a subspecies of Q. rubiginosa. 
Though we have observed great variation in this shell, we have never seen any transi- 
tion forms between the two species. It is rare in the Cumberland and the shells 
are rather small, measuring about 45 mm. long, 43 mm. high, and 25.7 mm. in diameter. 
The epidermis is clothlike and finely striate. The flesh is orange, in which respect 
it approaches rubiginosa. 

An example procured at Linton, Ky., had a dorsal baroque, and the mantle con- 
tained 4 marginal distomid cysts, a parasite which is especially frequent in this species. 

a Nautilus. S Nautilus, vol. xxm, no. 9, Feb., 1910, p. 116. 



MUSSELS OF CUMBERLAND EIVEE AND TRIBUTARIES. 61 

Where the pigtoe is found in abundance, as in some parts of the upper Mississippi, 
it is used quite extensively in the manufacture of buttons. It yields only a few- 
blanks per shell, however, and would not be a desirable species to propagate. 

71. Quadrula obliqua (Lemarck). Ohio River pigtoe. 

This is the most abundant, and, on this account, the most important, commercial 
species in the river, especially in the central portion, where it greatly exceeds any 
other species in number. 

The Ohio River pigtoe is a very good button shell. It is inferior to the niggerhead, 
both in luster and form, the sulcus on the side and the thinning out at the tip making 
it of unequal thickness; but, with the exception of the niggerhead, it is one of the 
best species. 

It is a rather prolific breeder. We found more gravid specimens of this than of 
any other species. The height of the spawning season is during the latter half of 
May and the earlier half of June. Occasional examples, however, may be found 
during the entire summer. Of five examined at Beasleys Shoals August 9, four 
were gravid. The portion of the gills used as marsupia varies greatly in different 
examples; it may depend upon the amount of ova fertilized and upon the age of 
the mussel. In some of the mussels the lower half of the outer gills are filled; in 
other cases the entire outer gills and quite frequently all four gills. Occasionally 
three gills, the two outer and one of the inner, contain eggs or young. There are 
no well-marked sulci between the conglutinates, which are rather thin and flat, 
resembling the seed of the green cucumber in general appearance. They are peculiar 
in that, when viewed from the side, they present a wavy appearance. This, so far 
as we know, is found only in the present species and enables one to distinguish the 
conglutinates even when found free from the animal. The wavy appearance is due 
to little pits in the anterior and posterior faces. A conglutinate of this species was 
found lying on the gravel bar in shallow water at Half Pone Bar June 16; the species 
was therefore spawning at that date. 

Dr. Ortmann has removed this species from the genus Quadrula and placed it in 
Pleurobema. All the examples he had examined up to that time had glochidia in 
the outer gills only. According to the data given above, its transfer to Pleurobema 
seems hardly advisable until the whole group is more thoroughly revised. 

72. Quadrula coccinea (Conrad). 

What appears to be an oblique form of Quadrula coccinea occurs rather frequently 
in the Big South Fork opposite Parkers Lake Station. Similar forms occur in the 
upper Cumberland down as far as Tear-coat Bar. In the main river these forms 
run into others in inextricable confusion, and nothing definite can be said about 
this speciee from the material at hand. 

Dr. Ortmann is of the opinion that Quadrula coccinea is a variety of Q. obliqua. 
In some of the northern rivers it seems to be a fairly constant and well-defined form. 

73. Quadrula solida (Lea). 

Only occasional. W 7 e obtained a few, principally at Indian Creek Bar. The 
shells were not typical and differed considerably from those found in the upper Mis- 
sissippi. The sulcus is very faint, and the nacre is not white but varies from pale 
rosy to purplish red. 

74. Quadrula plena (Lea). 

This appears to be a rare species in the Cumberland, and we obtained only a few 
scattered shells. They are all small and resemble very closely a much-shortened 
Q. obliqua, the compressed posterior portion being very short and the height of the 
shell being very great, considerably exceeding the length. The nacre is pale rosy. 

Mr. Boepple obtained this species in 1910 from Fords Island down to Martinsburg 
in the upper part of the river. 



62 MUSSELS OF CUMBERLAND EIVEE AND TRIBUTARIES. 

75. Quadrula pyramidata (Lea). 

Rare; we obtained a few examples in the vicinity of Mill Springs Bar. Our speci- 
mens have a broad furrow on the posterior half of the shell and differ from Q. obliqua, 
which they otherwise much resemble, by the umbones projecting far forward. They 
agree quite closely with Conrad's figure and description of Unio mytilcrides which 
Simpson regards as a synonym, except that the epidermis of our shells is black rather 
than brown and umbones are badly eroded. 

This is a very perplexing species. The extreme form, which, if it were only con- 
stant, would represent a very well marked and easily recognizable species, resembles 
an immensely overgrown Pleurobema clava in general appearance. Such specimens 
are rare; we have a few in the Washington collection. Our shells represent a sort 
of intermediate form between that and Quadrula obliqua. 

Mr. Boepple obtained examples from several stations in the upper river, to which 
portion it is apparently pretty well confined. 

76. Quadrula subrotunda (Lea). 

The young of this species have a general resemblance to Quadrula ebena, the 
niggerhead, but can be distinguished by their polished epidermis and broken rays 
near the umbones. We obtained only a few examples of these easily recognized 
shells. 

What is probably the adult of this species is occasional through the length of the 
river. We have not been able satisfactorily to connect the small shells with the large 
ones through a perfectly unbroken series, but up to the present can think of no better 
disposition to make of them. They have a black epidermis, with the umbones 
generally more or less eroded, and very much resemble an elongated ebena. These 
large shells are fairly common in the upper stretches of the river. A peculiarity of the 
old mussel is the rich orange color of the soft parts. At the blank factory at Clarksville 
they are known as the "long solid" and are regarded as one of the best button species 
of the river. None were found gravid. If they were to prove amenable to propa- 
gation, they might be profitable to plant in the upper part of the river and in s imi lar 
situations where ebena would not thrive. 

77. Quadrula ebena (Lea). Niggerhead. 

This important commercial species, which is generally regarded as the producer 
of the most valuable shell for the manufacture of buttons, is absent in the upper Cum- 
berland, and is abundant enough to be of considerable commercial importance only in 
the lower stretches of the river. 

The niggerhead is a deep-water shell and is rarely found in small rivers, or in such 
mussel beds as are found in shallow water. It seems in general to prefer mud to sand 
and gravel, and the percentage collected depends much upon the methods of collect- 
ing. Work in deep water will bring to light a larger percentage than wading or gath- 
ering by hand or a rake. 

The breeding season in the Cumberland begins in May and extends through the 
greater part of June, perhaps longer. In this species the condition of the development 
of the young can be roughly estimated by the appearance of the gill. When the ova 
pass down into the gill they are at first red, or carmine, probably because of an abund- 
ance of food material; as the glochidia develop they gradually fade out until the gills 
of a fully ripe niggerhead are of a dirty white color. 

There is not much variation in shape among the shells, some being elongate and 
others more rounded than the average. The shells show very little erosion, and the 
young exhibit the peculiar white patch near the umbone, as has been fully described 
by Lea. The nacre is rather frequently stained brown, and nearly all lack uniformity 
in thickness, the shell thinning out somewhat abruptly a little behind the middle of 

o Conrad, Monograph, p. 41, pi. xx. 



MUSSELS OF CUMBERLAND RIVER AND TRIBUTARIES. 63 

the ventral margin, leaving thin tips. The shell is easily distinguished from any 
other species in the river except from old examples of Q. subrotunda, which are always 
more elongate and always have yellow flesh. 

Q. ebena would probably thrive only in the lower parts of the river, although when 
the propagation of this species becomes feasible it may be worth trying in the upper 
river. 

78. Quadrula tuberculata Rafinesque. Purple warty-back. 

A careful study of our material, as well as of the evidence at hand from the literature, 
convinces us that Q. granifera and Q. tuberculata, though quite markedly distinct in 
typical cases, are really connected by intermediate forms. In some rivers, like the 
Tippecanoe at Delong, Ind., only strongly marked tuberculata are found. In others, 
like the Mississippi about Fairport, Iowa, only well-marked granifera are found. 
In such streams or portions of streams as contain both species they are indistinguish- 
able, or so connected by intergrades that no clear line of demarcation can be drawn 
between them. In the Cumberland, the first shells seen, in the lower part of the 
river, were identified provisionally as granifera; as we ascended the river some doubts 
as to the species began to appear, while in the upper tributaries the shells were pretty 
clearly identified as tuberculata. This naturally introduces the question as to influence 
of environment on shell form, which may be touched upon briefly here. 

The most striking and essential difference between tuberculata and granifera is one 
of degree of inflation, tuberculata being a flat form and granifera much inflated. We 
have a number of cases among the Unionidse where two otherwise similar shells are 
distinguished by this feature; among these are: Q. plicata, inflated, Q. undulata, 
compressed; D. dromas, inflated, D. caperatus, compressed. From our experience 
we are inclined to believe that one usually finds the compressed species in small 
streams, while the more inflated forms are found in large rivers. Often when a main 
stream has plicata, the little tributaries will have undulata, especially if they are 
rather shallow and swift streams with gravel bottoms. The more compressed form is 
better adapted to plow into the gravel or crawl under rocks and hold its position in a 
swift current, where the inflated form would present too much surface to the force of 
the water. In the softer mud and weaker current of larger streams an inflated form 
would be advantageous, helping to buoy up the animal. 

To state the situation precisely as we have found it, if one takes one of the larger 
rivers from source to mouth, and finds both tuberculata and granifera or plicata and 
undulata in the stream, the compressed form is likely to be in the upper stretches of 
the river while it is a small swift stream, and the more inflated form farther down in 
the main body of the river where the bottom contains more mud and the current is 
slower. Extreme forms of either species, so far as we know, are never found in the 
same bed, but where both are represented the forms run together. 

The literature relating to granifera and tuberculata is exceedingly interesting, but 
too long to give in detail. To understand the present status of the group, however, it 
is necessary to state that Simpson in his Synopsis removed these two species from the 
Quadrula pustulosa group, where they had been previously placed, making of them 
the subgenus Rotundaria on the basis of a "well-developed sulcus on the posterior 
slope and remarkable beak sculpture." The beak sculpture is well marked on tuber- 
culata but not so well, or almost absent, on granifera. Ortmann, finding only the outer 
gills used as marsupia in tuberculata, raised Rotundaria to generic rank. We have 
usually found only the outer gills of granifera at Fairport marsupial, although we have 
a record of one example with marsupia in all four gills. 

The species does not reach a very large size in the Cumberland. On account of 
its purple nacre it is of no value for buttons. 

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